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Sium 1 Apiaceae Heracleum L.1 Linnaeus, Sp. pl.: 249 (1753). Hemicryptophytes. Plant bristly at least on stems and petioles. Leaves pinnate, with a broad, inflated sheath. Flowers ± zygomorphic (petals white) or not (petals greenish yellow) 2. Fruit distinctly dorsiventrally flattened; carpophore filiform, divided; ridges straw-coloured to brown-yellow, 2 lateral wing-like and 3 low central ones (width of wings measured from the lateral veins3); vittae from the apex of the fruit, 4 clavate to saccate, red-brown (usually 4 dorsal ones c. 45–75 % as long as the mericarp and (0–)2(-4) shorter ventral ones). 1Heracleum Från Lars Fröberg 060103, och på slutet en äldre, redigerad fil (vet inte om alla ändringar beaktats av Lars)./MA061116 2 Föreslår omskrivning: Flowers ± zygomorphic (petals white) or not (petals greenish yellow) OK med denna /LF eller Flowers greenish-yellow (petals equal in size) or white (petals peripheral in the umbels enlarged, radiating) “Radiating” bör inte användas för “enlarged” utan för det att de ligger radiärt I förhållande till flockens mitt. Tror jag. Radiate/radiant har dock använts i ett flertal floror (se kommentar om detta under Caucalis) /LF Jag vet – synd att vi inte fick det i termlistan, hade man där skrivit nbågot i stil med ”används ibland i Apiaceae om kronblad som är större och radiära relativt flocken” (eller hur det nu var) så hade väl saken varit klar. Vi skulle kunna göra så i inledningen till umbellaterna - i princip bra metod men inledningen är så lång redan så jag tänkte inte på det utan försökte bara undvika en term till. – På korgblommiga har jag inte reflekterat så mycket över ordet, men det är något med konstruktionen som är känsligt och annorlunda hos umbellater; det är ju frågan om ”radiate/radiant” relativt hela den sammansatta flocken, inte den enskilda blomman (hos flockar som är ”radiate” är väl tvärtom den enskilda blomman ibland inte alls radiärsymmetrisk?), kanske inte ens den enskilda småflocken (? olika hos olika arter/släkten som har olika långt driven pollinatörsanpassning?). Man misstolkar lätt eller blir förvirrad, jag gjorde det – läshinder. (Vi skriver ju i stor utsträckning för dem som inte har /MA 3 förstår inte Det går en nerv precis vid varje vinges infästning till frukten, från vilken jag utgått när jag mätt vingarnas bredd /LF Är detta genomgående i alla umbellatsläkten eller i Heracleum eller i sphondylium? Om det är bara här bör det stå (measured from the lateral veins), är det genomgående i släkte eller familj bör det stå i inledning till släkte eller familj./MA Genomgående i Heracleum (inlagt kommentar i inledningen till släktet) /LF 4 OBS SE LÄNGST NED I DENNA KOMMENTAR! Följande ska läggas till om det inte framgår tillräckligt tydligt av (framdeles ändrad) bildtext eller annorledes: on the surface, from the apex of the fruit, lite omskrivet, hoppas det blev rätt jag vill stryka “on the surface” (vilket är det vanligaste hos umbellaterna, och är väl rätt självklart eftersom de syns på ytan?), samt “from the apex of the fruit” vilket dock kanske borde nämnas i Apiacéeinledningen eftersom det gäller generellt inom familjen (om det inte redan är gjort) /LF Om man (utan att den texten blir oläsligare lång) kan få redovisat dessa saker i inledningen kanske jag i sak är med på noterna (=ovanstående strykningar), men jag tänkte helt annorlunda: Jag försökte skriva den förklaring jag själv hade behövt. Jag förstod inte/var inte alls säker på/ att de där klunsarna på bilderna motsvarade de små prickarna i tvärsnittsbilderna. Jag hade heller inte anledning att anta annat än att vittae normalt går längsmed hela frukten, eftersom det inte står var tvärsnittet är lagt. Skulle kanske kunna avhjälpas med förklaring i figurtexten? /LF Hade för mig att vi nu enats om ventral/dorsal för umbellaternas delfrukter. Vill gärna ha det; vi har använt dem tidigare om blomdelar och sånt. Thomas brukar säga att adaxial/abaxial är så lika så de bör undvikas där de inte är omistliga – här är det väl tydligt nog vad som måste menas med ”rygg” och ”mage”? Dessutom står det ”dorsiventrally flattened”… Sium 2 Chromosome base number x=11; only diploids in Norden. Taxonomy. The classification of Nordic Heracleum (apart from H. sphondylium) has hitherto been very uncertain. There are 19th century records from Norden under several different names (e.g. H. giganteum Fisch. ex Horn., H. laciniatum Horn., H. palmatum Baumg., H. panaces L. and H. villosum. Fisch.); however, all these names have now been shown to be inapplicable: type material of H. giganteum, H. laciniatum and H. villosum is probably lacking, and H. palmatum and H. panaces are probably related to (or conspecific with) H. sphondylium subsp. sphondylium. In the late 20th century, the Heracleum taxon commonest in southern Norden was identified as H. mantegazzianum; the other fairly frequent taxon, with a more northern distribution, was referred to H. laciniatum or H. persicum (see further under H. persicum). There are also records from D of H. pubescens (see Variation under H. mantegazzianum).5 The four Nordic Heracleum species accepted here belong to three different groups. H. sphondylium belongs to section Euheracleum DC., H. mantegazzianum Somm. & Lev. and H. persicum Desf. ex Fisch. belong to section Pubescentia Mand., and H. platytaenium Boiss. belongs to section Villosa Mand. – See further under Pastinaca. i The giant species. Several of the impressive large Heracleum taxa were introduced to gardens, and escaped from cultivation already in the early 19th century; today, giant Heracleums are established and increasing in most of their distribution range in Norden: D scattered throughout, but probably more common in the eastern parts. N distributed in most of the lowlands; scattered in the southeastern parts, north to Op Øyer, and along the coast and in the fjords to Tr Tromsø (locally common around Oslofjorden and the surroundings of Trondheim and Tromsø). S scattered mainly in the southern parts, north to southern Vrm and southeastern Dlr; ÅsL Vilhelmina and PL Arvidsjaur; elsewhere casual (locally common in Sk and the surroundings of Stockholm). F fairly rare in the southern parts, north to central EH; casual further north to PeP Rovaniemi and Ks Kuusamo. Heracleum mantegazzianum and H. persicum are both rather frequent. Intermediates also occur, as well as some deviating morphotypes; it is possible that several genotypes have been introduced to Europe, subsequently intermingling (not least in botanical gardens), giving rise to the large variation that can be observed in Norden today. The taxa are not sympatric in their original distribution areas; the frequency of intermediates occurring where they meet (reported under various names, e.g. by Lid & Lid (2005) as H. mantegazzianum × tromsoensis), suggests that they hybridize easily and might be better treated as subspecies (but see further under H. persicum, Taxonomy). In any case, a thorough evaluation of all taxa is needed, including material both from their original sites and from areas where they are introduced. This treatment was based mainly on existing herbarium material, which is largely very incomplete; since several diagnostic characters are inavailable even in good herbarium specimens, also living material must be studied to understand the variation. 678A neighbour-joining analysis of molecular data (AFLP), made on material of giant Heracleum collected by local Rätta mig om jag har fel, om jag har argumenterat övertygande för adaxial/abaxial i fruktfallet! Jag trodde det var ad/abaxial som gällde – tidigare argument för dessa begrepp var att de är oberoende av ställningen av det aktuella organet, samt om det vridit sig eller ej. Jag är dock öppen för alternativa termer och tycker att dorsal/ventral är OK för delfrukterna /LF Mitt gamla förslag med ditt tillägg (första röda frasen kan strykas om det ersätts I inledningen): Fruit distinctly dorsiventrally flattened; carpophore filiform, divided; ridges straw-coloured to brown-yellow, 2 lateral wing-like and 3 low central ones (width of wings measeured from the lateral veins); vittae on the surface, from the apex of the fruit, clavate to saccate, red-brown (usually 4 dorsal ones c. 45–75 % as long as the mericarp and (0–)2(-4) shorter ventral ones) OBS om man skriver här att de är “from the apex of the fruit” så förenklar man och har en beredskap när man kommer till resonemangen under H. persicum!! OK, justerat så /LF 5 Material of Hornemann will be searched for in København by the author Har ej hunnit, kanske senare i vinter?/LF Senare i vår? /LF Jag har gått igenom alla Heracleum i det generella herbariet i Köpenhamn (utom H. sphondylium), och hittade en del Hornemann-material, men hittade inget material av H. laciniatum. Däremot fann jag H. giganteum, som han också beskrev. Materialet förefaller stämma med H. platytaenium, men arket var annoterad “1829” vilket var 10 år efter beskrivningen av H. giganteum, och dessutom avviker det något från beskrivningen i protologen; jag bedömer det sålunda inte som typmaterial /LF Kommentar om uppgiften av H. sosnowskyi från Danmark inlagd /LF 6 … den molekylära studien av de stora arterna… ger mig … gråa hår och jag vet inte vad jag ska tro om resultaten det hela ser snyggt ut med tre grupper: 1 = H. mantegazzianum, 2 = H. sosnowskyi och 3 = H. persicum. Jag kan själv bara utgå från morfologin och har endast sett i princip nordiskt material som ingått i studien (delvis samlat av mig Sium 3 själv); materialet som ingår i H. persicum-gruppen är morfologiskt heterogent, men det hindrar ju inte att de kan vara molekylärt lika; materialet från Danmark i H. sosnowskyi-gruppen är lite avvikande och kan ju motsvara H. sosnowskyi, men jag har tyvärr inte förstått mig på denna art. Sen är det analys-metoden: jag tror att AFLP (amplified fragment length polymorphism) används för att analysera inomartsvariation, och om jag fattat metoden rätt bör den vara betydligt osäkrare för artklassificeringar än sekvensering - men jag är inte säker. Vore kanske värt att kolla med någon som är bevandrad i DNA-analyser. Jag är tacksam för alla synpunkter på avsnittet. Meddela mig om du vill se texten och dendrogrammet på deras analys, så kan jag skicka kopia på det. (mail LF >MA 081027) Metodologiskt är analys av AFLP data en metod där man utifrån ett totalDNA klipper upp det i småbitar med restriktionsenzymer. De erhållna bitarna körs med elektrofores på en gel och ger ett bandmönster som kan liknas vid en streckkod eller "fingerprint". De olika fragmenten kodas som finns-finns inte och det genererar till slut en datamatris som kan köras i ett analysprogram. Så långt är allt väl. Fragmentanalys är n bra metod för studier av populationsvariation och i taxa med hybrider eller i övrigt närstående komplex av taxa. Emellertid är Neighbor-joining en metod som är starkt ifrågasatt. Ett av skälen är att man alltid erhåller ett enda träd som resultat och till skillnad från metoder som bootstrapping och jackknifing erhåller man ingen test av stabiliteten i de olika grupper man ser i det erhållna trädet. Jag skulle personligen anse att grupper som erhållits i ett neighbor-joining träd skulle anses suspekta till dess man visste om de hade gott bootstrap eller jackknifesupport Arne (mail 081027) …stycket som refererar studien är lite för detaljerat när man ändå inte kan lita på den och skulle föreslå något i stil med följande: "A neighbour-joining analysis of molecular data (AFPL), made on material of giant Heracleum collected by local botanists in Europe and western Asia, was presented in Jahodova et al. (2007). The results showed an assemblage of the material in three groups, tentatively were referred to H. mantegazzianum, H. persicum and H. sosnowskyi (based on the determinations made by the collectors). Unfortunately, it is not possible to make any general taxonomic conclusions on the study, since the molecular data cannot be connected with morphological information (due to absence of vouchers in many cases). A specimen from D Sjæ København was reported as H. sosnowskyi as new for Denmark based on this study. However, the specimen is morphologically intermediate between H. mantegazzianum and H. persicum, and the the true status of the name H. sosnowskyi sensu Mandenova is unclear." Thomas (mail 081027) 7 Faurholdt & Hinke (2007) reported H. sosnowskyi as new to Denmark based on a specimen from D Sjæ København in this study; however, the specimen is morphologically intermediate between H. mantegazzianum and H. persicum, and the identity of H. sosnowskyi sensu Mandenova needs to be determined. Men det var bara en omformulering - jag förstår inte riktigt. Du säger samma sak som tidigare, kommenterar inte AFLP-resultaten?– avviker detta ex från övriga I sosnowskyi-gruppen, eller är slutsatsen att alla kan misstänkas vara såna intermediärer? Nej, det är fullt möjligt att det danska morfologiskt stämmer bra med övriga i gruppen. Problemen är dels att vouchers verkar saknas för de flesta övriga, samt att identiteten av H. sosnowskyi sensu Mandenova verkar oklar. Jag lånade material av H. sosnowskyi från Tartu, men det var heterogent och föga överensstämmande med Mandenovas beskrivning – det är möjligt att den som samlade materialet i Kaukasus har koll på arten, men det finns också en risk att materialet bestämdes utifrån lokalerna (Armenien där arten är uppgiven). FÖRSÖK TILL OMSKRIVNING: “...based on a specimen from D Sjæ København in this study; the specimen was redetermined to H. sosnowskyi based on the molecular results. However, morphological data should also be consulted when assessing the identity of a specimen in such a complex group; in the present account it is regarded as an intermediate between H. persicum and H. mantegazzianum, although with the leaves having broader and more rounded lobes. Furthermore, since H. sosnowskyi is closely related to H. mantegazzianum, its identity must be established by comparison with original material, and its distinctness against H. mantegazzianum should also be assessed” /LF Sium 4 botanists in Europe and western Asia, was presented in Jahodova et al. (2007). The results showed an assemblage of the material in three groups, tentatively referred to H. mantegazzianum, H. persicum and H. sosnowskyi. Unfortunately, no general taxonomic conclusions can be drawn from the study, due to method problems (inadequacies regarding vouchers and statistical testing). Based on a specimen included in that study, H. sosnowskyi was reported from D Sjæ København (Faurholdt & Hinke 2007); however, the correspondence of the Danish specimen to Mandenova’s H. sosnowskyi (a ii name with unclear status ) could not be verified, and it is here treated together with the many other Nordic specimens morphologically intermediate between H. mantegazzianum and H. persicum. Diagnostic characters. Important features for determination of Heracleum specimens are life form, leaf-shape and indumentum, as well as indumentum on rays, pedicels and ovaries/fruits, and width of vittae. Many specimens lack information on ”gross morphology” and are incomplete, consisting of only very small parts of leaves and umbels in flowering stage, leaving type of indumentum on different parts of the plant as the main character that can be studied. Indumentum on rays and pedicels may consist of papillae, hairs or both in varying densities. They are usually better developed on the rays than on the pedicels. The hairs are either glandular (i.e. translucent and flexuose) or not glandular (whitish and ± stiff), but intermediate types may also occur. The papillae 8 Det gröna stycket Furthermore, some specimens from D, N and F were located in the H. persicum group. The vouchers seen were morphologically somewhat heterogeneous: a specimen from F V Karkkila corresponded fairly well with H. persicum, while two specimens from D Sjæ Roskilde and another one from N Tr Balfjord deviated by having shorter and fewer hairs on the ovaries/fruits and slightly smaller papillae on the rays and pedicels, and they are here referred to intermediates between H. persicum and H. mantegazzianum. Unfortunately, it is not possible to make any general conclusions on the study, since it is based only on molecular data. är oklart. Finns det vouchers för de specimens du nämner nu (grön markering)? Jag har sett vouchers av den danska, en norsk och en finsk /LF Avser tredje meningen att just dessa vouchers är dåliga, eller att ett eller flera av just dessa specimens saknar vouchers, eller syftar du på undersökningen i dess helhet i denna mening? Nej, det avser ‘gross morphology’ som hör hemma under ‘diagnostic characters’ – kom med av misstag här /LF Sista meningen: Fattar jag rätt att undersökningen är dålig såtillvida att man inte har vouchers till allt material – eller är det fler grundläggande brister? Är det inget annat kan man säga det kortare. Nej, det brister även i sättet att analysera data: Man har fått in material av blad på vilka man gör en analys av DNA och får ett dendrogram. De kollekter som hamnar ‘fel’ utifrån ursprungsbestämningen får byta namn utan att vouchers kollas. Jag är även lite osäker över relevansen i typen av DNA-analys (jag tror AFLP innebär att man med restriktionsenzymer erhåller DNA-bitar som körs ut på en gel och ger ett bandmönster vilket analyseras – jag är osäker på detta, men om det stämmer så är det en betydligt osäkrare analysmetod än sekvensering) /LF Hur mycket materail, hur stor andel saknar vouchers? Jag har gjort förfrågan på vouchers och fick några napp, men tyvärr inte på de grupper jag var mest intresserad av. Dock har jag som sagt sett en del av det nordiska /LF Allmänt om texten om den här studien: Nu går liksom kritiken mot undersökningen ”parallellt” med dina påståenden om materialet – de möts aldrig riktigt. Du har ju skrivit att de ”tentatively” kallar saker enligt bestämmarnas namn – det skulle kunna betyda att man fått fin korrelation mellan morfologi och molekylärdata, men det skulle också kunna betyda att man försökt bevisa det man ville, med ett alldeles för litet material. Du är för ospecifik också. Det är för många okända i ekvationen! Jag håller med om att jag formulerat mig luddigt och med dåligt koppling till det nordiska. Jag ger nytt fylligare förslag FÖRSÖK TILL OMSKRIVNING: “Furthermore, some specimens from D, N and F were located in the H. persicum group. The vouchers seen were morphologically somewhat heterogeneous: a specimen from F V Karkkila corresponded fairly well with H. persicum, while two specimens from D Sjæ Roskilde and another one from N Tr Balfjord deviated by having shorter and fewer hairs on the ovaries/fruits and slightly smaller papillae on the rays and pedicels, and they are here referred to intermediates between H. persicum and H. mantegazzianum. Unfortunately, it is not possible to make any general conclusions on the study, since it is based only on molecular data. /LF VARIANT FRÅN THOMAS: “A neighbour-joining analysis of molecular data (AFPL), made on material of giant Heracleum collected by local botanists in Europe and western Asia, was presented in Jahodova et al. (2007). The results showed an assemblage of the material in three groups, tentatively referred to H. mantegazzianum, H. persicum and H. sosnowskyi (based on the determinations made by the collectors). Unfortunately, it is not possible to make any general taxonomic conclusions on the study, since the molecular data cannot be connected with morphological information (due to absence of vouchers in many cases). A specimen from D Sjæ København was reported as H. sosnowskyi as new for Denmark based on this study. However, the specimen is morphologically intermediate between H. mantegazzianum and H. persicum, and the true status of the name H. sosnowskyi sensu Mandenova is unclear.” OK med mig /LF Sium 5 varies in shape and may be translucent or whitish. Similar types of indumentum also occur on the ovaries/fruits. Length of indumentum on ovaries/fruits is rather constant in time, but vittae should be measured on ± ripe fruits. A specimen should include at least the following: an umbel or some of the largest umbellules (preferably from the primary umbel) in flower, fruits that are as ripe as possible, and a piece of a lower leaf with rachis (preferably from the apical leaflet). Information on height, number of stems, pigmentation, leaf division and shape of umbels should be included if possible; a photograph may be very valuable. Heracleum stevenii Mand. was reported from S Upl Stockholm (Lindberg 1983), based on material intermediate between H. mantegazzianum and H. persicum, and from D Sjæ København (Madsen & Lyck 1991), based on material redetermined to H. platytaenium. 1 - Leaves bristly on both sides (upper side sparsely so, lower side with bristles longer than 0.2 mm); pollen grains shorter than 48 μm; dorsal vittae thin (median ones 0.15–0.25 mm wide, lateral ones 0.2–0.4 mm wide)................................................................................................................................ 1. H. sphondylium Leaves glabrous or puberulent at least beneath (upper side glabrous or sparsely bristly, lower side with up to 0.2 mm long hairs); pollen grains longer than 48 μm; dorsal vittae thick (median ones 0.4–1.2 mm wide, lateral ones 0.5–1.5 mm wide) ......................................................................................................................2 2 - Leaves with rounded lobes, puberulent beneath9 and with sparse bristles above ........... 4. H. platytaenium Leaves with acute lobes, glabrous above and glabrous or hairy (but not puberulent) beneath .....................3 3 Leaves with elongated, narrow lobes and teeth with concave sides; umbel rays with narrow, translucent, patent papillae and/or glandular hairs; ovaries/fruits with c. 0.5-1 mm long glandular hairs; dorsal vittae distinctly expanded10 ................................................................................................ 2. H. mantegazzianum Leaves with short, wide lobes and short teeth with ± convex sides; umbel rays with broadly triangular, whitish, ± ascending papillae, and usually also with non-glandular hairs; ovaries/fruits with whitish, nonglandular hairs usually longer than 1 mm; dorsal vittae slightly expanded ........................... 3. H. persicum - 1. Heracleum sphondylium L. Linnaeus, Sp. pl.: 249 (1753). – Type: Clifford Herbarium 103, Heracleum 1, fol. 3 (BM) lectotype, sel. by Reduron & Jarvis, Regnum Veg. 127: 53 (1993). F ukonputki. S björnloka. Biennial, to 1.4 m; taproot 7–16 mm thick. Stem solid, or secondarily hollow; basal part 4–9 mm thick, distinctly to indistinctly sulcate or angular, pale greenish or sometimes purplish, not or slightly glaucous, and usually densely bristly (sometimes with hairs or papillae interspersed); upper internodes distinctly to indistinctly sulcate, with a sparse to dense cover of papillae or hairs (rarely with a few bristles interspersed). Leaves 2–3(–4) at the base and 3–5(–6) on the stem (the innermost basal one or the lowest stem leaf is the largest); sheath broad, sometimes purplish; petiole 8– 27(–32) cm; blade 1-pinnate, (10–)15–31(–36) × 8.5–22 cm (length/width ratio 1.1–1.9), lower side usually rather densely covered with bristles, upper side sparsely covered with coarser bristles. Leaflets (1–)2(–3) pairs; angle leaflet/rachis (35–)45–70°. Apical leaflet 1–2-pinnatifid, with 1–3 pairs of lobes (sometimes elongated and narrow); petiolule 2–8 cm; blade 6–18 × 8–19 cm, with a length/width ratio of 0.7–1.1(–1.4); base usually cordate, sometimes iii truncate; lobe apices usually acute; margin crenate with usually purplish tips . Lateral leaflets 1–2-pinnatifid, with 1– 2(–4) pairs of lobes; petiolule 0.5–5 cm long, sometimes sessile; blade 5–17 × (3.5–)5–12(–15) mm (length/width ratio 0.8–1.7); base broadly and often obliquely cuneate to cordate. Umbels flat to slightly convex; peduncle 10–31 cm; rays straight or bent inwards, 5.1–10.5 cm, usually densely papillose on the adaxial side, with narrow, acute and translucent papillae often sparsely interspersed with short glandular hairs. Bracts 0(–7), persistent. Umbellules 8–24(–34); pedicels 0.6–1.5(–2.1) cm, usually densely papillose (sometimes also with sparse glandular hairs) on the adaxial side. Bractlets 2–10, persistent, 2–15 × 0.2–0.9(–1.2) mm, usually hairy, without a membranous margin. Flowers 24–38(–48) per umbellule; sepals 0–0.3 mm; petals variable in size and colour (see subspecies); filaments 2–4 mm; anthers 0.7–0.9 mm. Pollen grains 38–47 × 19–26 μm 9 Villous ska enl termlistanvara ”covered with long, weak hairs” – 0,2 mm låter mer som mycket korta hår, pubescent eller puberulent. (nod 2) Justerat /LF 10 Omskrivningsförsök: OK, bra – har dock svårt att finna ord för skillnaden i svällning: om man följer oljekanalerna nedåt på frukten, så sväller de mer plötsligt hos mantegazz., medan de hos persicum sväller mer gradvis /LF Blir de ungefär lika tjocka till sist så bör man väl säga ”abruptly” kontra ”gradually” och skippa ”distinctly” och ”slightly”./MA OK, det är främst tjockleken (bredden) som skiljer – jag låter distinctly/indistinctly stå kvar /LF Sium 6 (length/width ratio 1.6–2.2)11. Fruit obovate to elliptic in outline. Mericarps 5.8–9.5 × 4.5–6.8(–7.7) × 0.4–0.8(–1) mm (length/width ratio 1.1–1.6); wings 0.4–0.7 mm wide; dorsal vittae not expanded at the apex, median ones 3.5–5 × 0.15–0.25 mm, lateral ones 3–5 × 0.2–0.4 mm; stylopodium flattened, 1.1–1.6 mm wide; style 0.7–1.5(–2.2) mm, directed upwards to slightly outwards. – Mid-summer to late summer. 2n=22 (F V, U); 2n=22 + 0–3B (S Sk 5, Srm). [2n=22]12 Distribution.13 Nem–BNem. – Mainland Norden, resident in most lowland areas, casual in S Lappland below the mountain regions and northernmost F; subsp. sphondylium is distributed in the southern and southwestern parts (archaeophytic or perhaps native in D and western N), subsp. sibiricum has a more northern and eastern range and is much more common, and intermediates are frequent in eastern D, southern S and western N. – Map xxx. Europe except the Mediterranean and the northernmost parts, W Siberia and N Africa. Introduced in North America, South America and New Zealand. Habitat. Dry to moist, nutrient rich, mull-rich, clayey or sandy ground, in sun-exposed to shaded places. Usually in man-made sites, such as roadsides, railways, overgrowing fields, pastures and ruderal places, but also in seminatural sites, such as woodland margins, glades and seashores. Sometimes forming large populations, e.g. along roadsides and railways; perhaps favoured by fertilization. Biology. The seeds are probably mainly wind-spread. – Many specimens have a reduced pollen fertility, although their morphology does not indicate hybridization between the subspecies. Weimarck (1978) reported a reduced pollen fertility in H. sphondylium from various sites in Europe, which was later explained by translocation heterozygosity in the chromosomes (Weimarck et al. 1979). There are also reports of B-chromosomes in several of the studied populations, but no correlation was found between B-chromosomes and reduced fertility, neither on the male nor on the female side (Weimarck 1978). 11 struken text: All flowers of the primary and secondary umbels are bisexual, or the outer flowers of the primary umbels are female in both subspecies. The flowers of the tertiary umbels are either bisexual or male Jag är osäker på könsförhållandena, eftersom ståndarna kan ha ramlat av från de yttre blommorna; föredrar därför att stryka meningarna/LF 12 Uotila & Pellinen 1985; Weimarck 1978 (för Norden) /LF 13 Probably partly native (see subspecies). D common to scattered in most parts, but rare in western Jylland. N scattered to fairly rare north to central VFi (mostly in the lowlands; more common in the southeastern parts); rare in Øfi[OBS stämmer dåligt med nuvarande karta för Tr-Öfi. Men jfr föregående kommentar till hela distributionavsnittet för arten.]. S common to fairly common in the lowlands, north to Dlr (around Siljan), southeastern Jmt and southwestern Ång, further north scattered to rare mainly in the lowlands; casual in the mountains up to the forest belt. F fairly common to scattered north to KP and Kn, further north gradually rarer; casual in EnL[OBS casual även i InL, och här framgår inte skillnader i ålder. Men jfr föregående kommentar till hela distributionavsnittet för arten. OK, kompletterat /LF]. Om man har två distinkta underarter med olika ursprung blir den här sammanfattningen rätt meningslös. Försök från Magdalena.: Distribution and habitat. Nem–BNem. – Mainland Norden, resident in most lowland areas; subsp. sphondylium is southern and southwestern (archaeophytic or perhaps native in D and western N), subsp. sibiricum has a more northern and eastern range and is much more common, and intermediates are frequent in D, southern S and western N. OK, inlagt /LF lokaler som inte täcks in av någon av underarterna: N central VFi S common to fairly common in the lowlands, north to Dlr (around Siljan), southeastern Jmt and southwestern Ång, further north scattered to rare mainly in the lowlands; casual in the mountains up to the forest belt. - det bör istället/också stå noggrant för det som underartsbestämts. F casual in EnL. Sium 7 Variation. Heracleum sphondylium varies extensively, e.g., in size, leaf shape and division, colour and degree of radiation of petals, and in the occurrence and length of fruit indumentum. The species has been divided into 9 subspecies in Europe (Brummitt 1968). Two of these occur in Norden, viz subsp. sphondylium and subsp. sibiricum. Intermediates are common in areas where the subspecies are sympatric (see distribution) and they show a significant variation, combining the characters of the subspecies in different ways. The taxa have often been treated at specific level (e.g. Elven 2005, Hämet-Ahti et al. 1998), but are better regarded as subspecies since no reduced fertility has been found in intermediates. Specimens with papillose fruits occur both in subsp. sphondylium (= var. chaetocarpoides Gawlowska), subsp. sibiricum (= var. chaetocarpum Thell.) and intermediates, and the character is regarded as taxonomically insignificant. Similar taxa see Angelica archangelica, A. sylvestris, Levisticum officinale and Pastinaca sativa. 1A. subsp. sphondylium H. sphondylium subsp. australe (Hartm.) Ahlfv. (1901). – H. australe Hartm. (1846). D Almindelig Bjørneklo. F etelänukonputki. N kystbjønnkjeks. S vit björnloka. Apical leaflet 8–17 × 9–19 cm. Bractlets (2–)5–10, 5–15 × 0.3–0.9(–1.2) mm. Outer flowers of outer umbellules zygomorphic; petals white (sometimes with a violet tinge), 3–8.5 × 3.5–8.5 mm, bifid (apical cut 1.5–4 mm deep), usually with short hairs on the outer side. Fruit usually densely hairy, rarely papillose (indumentum sometimes destroyed during the drying of specimens). Distribution and habitat. Nem–BNem. – Resident mainly in southern Norden; archaeophytic or perhaps native in D and western N; elsewhere brought in later via harbours and railways, and probably also with grass seed and cereals. – D in most of the distribution area of the species in e.g. roadsides, pastures and forest margins; commonest in ØJy, southeastern NJy, FyL and LFM Lolland, less common in Sjæ. N scattered along the coast and the fjords in e.g. pastures, shrubby woodland and roadsides; Oslofjorden 7 localities, and from VA Farsund to SF14 Førde; casual in Te Kragerø, Tokke (inland locality on roadside), and NT Grong 1995 (railway staiton). S Sk 15 localities (e.g. riversides, roadsides and parks) in the western part, close to the localities in D Sjæ15, Bl Karlshamn known since 1871 (railway and roadside), Klm Kalmar since 1872 (e.g. railway), Ukna since 1952 (roadside), SmI Sävsjö since 1961 (railway), Hl Falkenberg 1921, 1951, BhG Göteborg since 1922 (waste ground), Srm Hölö (Tullgarn, park and roadside since 1864); casual north to Ång Örnsköldsvik 1989 (railway). F U Helsinki 5 localities (e.g. roadside and woodland, since 1932), EP Vaasa since 1945 (dock and railway), Kn Hyrynsalmi since 1945 (with german troups); casual north to PeP Tornio 1985, and Ks Kuusamo 1969 (fallow field). – Map xxx. W and C Europe (e.g. the Alps), N Africa. 1B. subsp. sibiricum (L.) Simonk. Simonkai, Enum. Fl. Transs.: 266 (1887). – H. sibiricum L. (1753). – Type Linnaean Herbarium 352.5 (LINN) lectotype, sel. by Reduron, Nordic J. Bot. 22: 84 (2002). H. sphondylium subsp. flavescens (Willd.) Soó (1973). – H. flavescens Willd. (1809). D Grønblomstret Bjørneklo. F idänukonputki. N sibirbjønnkjeks. S sibirisk björnloka. Apical leaflet 6–14 × 7.5–12 cm. Bractlets (0–)3–7, 2–7 × 0.2–0.5(–0.7) mm. Outer flowers of outer umbellules not or slightly zygomorphic; petals green-brown or brown-yellow (sometimes with a violet tinge), 1.7–2.6 × 1.1–2.4 mm, entire, glabrous. Fruit glabrous or sparsely to rather densely papillose, never hairy. 16 Distribution and habitat. Resident in most parts of Norden, but not native in large areas of D and F (for habitats see the species). – D scattered in Sjæ (more common than subsp. sphondylium), rare in LFM and eastern Jylland. N fairly 14 struket: Askvoll and 15 Enligt Thomas oerhört markant skillnad inom landskapet, typ vikarians. Kan denna utbredning tänkas höra ihop med det danska? Som läsare undrar jag över att saken inte diskuteras. Jag kollade lokalerna i Skåne o. Sjælland och de ligger verkligen i nära anslutning – kompletterat /LF 16 Jfr under arten. S och F måste förstås ändras om texten på arten ändras, och helst i vilket fall som helst – man bör kunna få besked åtminstone om nord/syd/öst/västgränser och tyngdpunkter!. Justerat på arten; i S o. F bedömer jag att ssp. sibiricum är utbredd i ± hela artens utbredningsområde /LF Sium 8 common to scattered in the southeastern parts west to AA Bykle and north throughout the highlands of Op and western He; fairly rare in the western coast and the fjords from SF Leikanger and Vik to NNo Gildeskål; Tr Tromsø and Nordreisa, ØFi Sør-Varanger. S and F in most of the distribution area of the species. – Map xxx. N and E Europe, W Siberia. Variation. Specimens with narrow-lobed leaves and entire to coarsely crenulate margins (e.g. from S Sk, Öl, Srm and Upl) have been recognized as var. angustifolium auct., but no correlation was found with other characters. 2. Heracleum mantegazzianum Somm. & Lev. Sommier & Levier, Nuovo Giorn. Bot. Ital., nov. ser. 2: 79 (1895). – Described on cultivated material (from Caucasus). D Kæmpe-Bjørneklo. F kaukasianjättiputki. N kjempebjønnkjeks. S jätteloka. Literature. Tiley et al. 1996, Often & Graff 1994, Pysek et al. 2007. Hemicryptophyte (hapaxanthic).17 Plant to 2.2–3.2 m, usually with only one stem; scent sharp, aromatic or scentless. Stem sparsely to densely hairy, hollow; basal part 50–100 mm thick, usually sulcate, with purplish patches and bristlelike hairs (towards the apex with softer hairs). Leaves18 with a rather broad, usually purplish sheath; petiole 40–90 cm; blade 1-pinnate (almost ternately divided), 80–130(–170) × 90–150 cm, with a length/width ratio of 0.7–1(–1.4), lower side densely to sparsely covered with short hairs or glabrous, upper side glabrous. Leaflets 1–2 pairs. Apical leaflet 2pinnatifid, with 3–4 pairs of primary lobes; petiolule 3–10 cm; blade 16–64 × (17–)23–72 cm (length/width ratio 0.7– 1.1); base truncate to deeply cordate; margin doubly dentate to doubly serrate, with acuminate teeth; lobe apices usually long, narrowly acute to acuminate. Umbels flat to slightly convex, 9–15 cm high and 25–60 cm wide;19 rays straight or bent inwards, 14–32 cm, on the adaxial side with translucent, patent papillae, and/or densely hairy, with long or short glandular hairs. Bracts (0–)10– 1820, persistent. Umbellules 60–120; pedicels 1–3.5 cm, on the adaxial side with papillae and hairs as in the rays (sometimes more densely hairy). Bractlets 10–1821, persistent, 7–15(–22) × 0.5–1.3 mm, sligthtly to distinctly hairy. Flowers (28–)34–73 per umbellule; sepals 0.3–0.9(–1.2) mm; petals 6–10.5 × 6–11(–13) mm (length/width ratio 0.6– 1.4), bifid (apical cut 3–7 mm deep), with 2–3.5 mm wide lobes and indistinct to distinct veins; filaments (2.5–)3.5–5 mm; anthers 0.9–1.4 mm. Pollen grains 50–74 × 28–42 μm (length/width ratio 1.3–2.4). Fruit obovate in outline, with a sparse to dense cover of c. 0.5–1 mm long glandular hairs, sometimes also with minute, upwards directed hooks on the lateral parts. Mericarps 8.5–15 × 5.5–9.5 × 0.6–1.4 mm, with a length/width ratio of 1.2–1.7(–2); wings 0.6–1.2(– 1.5) mm wide; dorsal vittae distinctly expanded at the apex, the median ones 5–9.5 × 0.7–1.2 mm, the lateral ones 4.5– 8.5 × 0.8–1.5 mm; stylopodium flattened, 1.2–2.3 mm wide; style 1.8–3.7 mm, directed ± outwards. – Early summer to mid-summer. Distribution. Heracleum mantegazzianum was introduced as an ornamental; the first verified record of the species as an escape is from 1903 (S Srm Nynäshamn). The species is probably more common than shown here (only records verified by vouchers have been included). – 22D a recent introduction, but probably widespread; eastern Sjæ 5 localities 17 Monocarpic avser väl att den växer i flera år innan den blommar en gång, sen dör den? Vi har inte använt ”monocarpic” tidigare, men ”short-lived hapaxanth” för såna som inte är strikt bienna – så ”hapaxanth”, gärna med ”long-lived” eller ”short-lived”, skulle vara bättre – om det nu är samma sak som du menar. (Perennial avser ibland, och vi har väl i stort sett använt det så, flerårig med återkommande blomning.) OK, justerat/LF Ska det inte vara long-lived? Hur gammal blir den? Jo, den lever 2-7 år; föreslår “short- to long-lived hapaxanth” /LF 18 antal står på sphondylium saknar uppgift (mindre viktig karaktär) /LF 19 peduncle? Saknar uppgifter /LF 20 Obs, antal svepeblad från fältmätningar istället för herbariekollekter/LF 21 Obs, antal svepeblad från fältmätningar istället för herbariekollekter/LF 22 Lite ändrat, OK? Jfr kommentar på Sverige. Justerat (endast tillfällig i VJy) /LF Om allt utom Vjy avsåg bofasta förekomster så får man ändra; som det stod trodde jag att allt var tillfälliga Menar du uttryckligen ”localities” eller kan man skriva “records”? ”localities since + årtal” kan tolkas både som 5 fynd sedan det första, och (lite mer osannolikt, men logiskt inte omöjligt) som 5 lokaler som alla bestått sedan det året. Förra versionen: Sium 9 (first record 1953), FyL 3 localities (first record1951), NJy Hassing 1953, Rolds skov 1980; VJy Søndre Felding 1962, Ulfsborg 1968 (loading place). N a recent introduction (first record from Ho Fana 1958), resident at least at Oslofjorden; casual in coastal regions north to NT Leksvik. S established in the south (northernmost record Dlr Krylbo (roadside, since 1960), mainly in coastal regions; Gst Torsåker 2006, Mpd Njurunda 199023. F casual north to PeP Rovaniemi 1986 (park) and Ks Kuusamo 1998 (roadside)24. – Map xxx. Caucasus. Habitat. Sun-exposed to rather shaded, rather moist, nutrient-rich habitats; indifferent to soil pH. Common in manmade sites, such as road-sides, construction areas, landfills, railway yards, dumps, gardens and parks; frequently also in seminatural sites such as deciduous and mixed woodland, unmanaged grassland 25, seashores, lakeshores and along rivers. At several localities ( e.g., in S Sk and Srm/Upl Stockholm are) forming dense and large stands. Biology. Heracleum mantegazzianum is mainly hapaxanthic, and the age of flowering individuals ranges from 2–7 years. It is andromonoecious, self-compatible and protandrous; among the pollinators are coleopterans, dipterans, hemipterans and hymenopterans. The seeds are spread by wind (mainly over short distances), water and human activities (Moravcová et al. 2007).26 The germination rate is very high under favourable conditions; the seed bank is not long-persistent. The species contains furanocoumarins which, together with UV radiation, cause serious skin injuries (photodermatitis; Hattendorf et al. 2007). Mechanical and chemical methods are used to control the species, but grazing by sheep is considered the most effective means. Coleopterans (mainly curculinoids), dipterans, hemipterans (mainly aphids) and lepidopterans are the most important of the invertebrates feeding on H. mantegazzianum (Hansen et al. 2007). In its native area, H. mantegazzianum is the target of several parasitic fungi, but few of them occur in the D a recent introduction; eastern Sjæ 5 localities since 1953, FyL 3 localities since 1951, NJy Hassing 1953, Rolds skov 1980; casual in VJy Ulfsborg 1968 (loading place) and Söndre Felding 1962; probably widespread. OBS som standard skriver vi inte ut ”casual” – ”casual” är underförstått när det efter semikolon kommer lokaler med årtal. Det bör alltså inte stå ”casual” före VJy, men om det är/antas vara/ bofasta förekomster i övrigt så ska det justeras där! OK justerat /LF 23 Förut: “resident in coastal regions north to Dlr (Krylbo on a roadside since 1960); casual north to Mpd Njurunda 1990” Obs Krylbo ligger ej i coastal regions… Måste skrivas om. Det finns inte heller några andra tillfälliga förekomster redovisade än Njurunda, så ”north to” är tokigt. Såhär kanske: established in the south (northernmost record Dlr Krylbo (roadside, since 1960), mainly in coastal regions; casual in Gst Torsåker 2006 and Mpd Njurunda 1990. OK med liten justering /LF Vi har genomgående tillfälliga förekomster på slutet efter semikolon, utan särskilt påpekande att de är tillfälliga, och här kan det knappast missförstås./MA 24 ”casual north to” + årtal funkar EJ. RAINO could you summarize with the help of the documentation? FÖRSLAG: “F V Lohja 1986, EH 7 localities since 1975, PS Kirkonsalmi 1976, Iisalmi 1977, EP Vaasa 1988, KP Reitsjärvi 2000, Raahe 1989, OP Oulu 1997, 1998 (old lawn), PeP Rovaniemi 1986 (park), Ks Kuusamo 1998 (roadside).” /LF 25 =igenväxande slåttermark, eller igenväxande betesmark, eller avses slåttermark som slås men ej efterbetas? Meadows brukar inte användas för betesmark så man blir förvirrad. Felskrivet; skall vara ‘ohävdad gräsmark’ - justerat /LF (ändrat ”ungrazed” till ”unmanaged” som bättre torde motsvara ”ohävdad”/MA) 26 Hurså? Belägg? Utdrag från Moravcová et al., Seed Germination, Dispersal and Seed Bank in Heracleum mantegazzianum (In ‘Ecology and Management of Giant Hogweed’, eds. Pysek et al. 2007: 74-91): “For plants of 2 m heigh, 60-90% of the fruits fall within a radius of 4 m. Dispersal by wind is argued to be important only over short distances. Fruits may float for 3 days (or if 6 months old for 8 hours). Such time is likely to be sufficient for spreading a long distance, especially by fast-flowing streams. Humans may spread fruits stuck to car tyres along roads, move them to new locations with soil transport, or deliberately transport decorative umbels with dry fruits. Given that longdistance dispersal is important to the success of possible invasion, dispersal by water and humans seem to be the most significant factors in this respect.” – Jag lägger in referens till det /LF Kanske vill man ge mer utförlig info – jag låter kommentaren stå kvar så länge /MA OK /LF OBS! Jag har strukit den generella litteraturhänvisningen till “Pysek et al. 2007”, eftersom jag lagt in referenserna från de olika kapitlena i denna specifikt istället /LF Sium 10 introduced range (Seier & Evans 2007). Variation. 27A deviating morphotype (leaves with no elongated lobes, umbels with fewer rays (9–40) and flowers with purplish anthers but without distinctly radiating peripheral petals) has been seen from D ØJy Viborg 1941 (garden relic), Odder 1971, VJy Skjern 1968 (ditch), Sjæ Gentofte 1961, Søndersø 1965 (river), Langholmen 1965, N 28Te Langesund 1951, Porsgrunn 2005, S Srm Nynäs 1903 and Upl Danderyd 1945, 1981 (railway station). This variant, which is also somewhat smaller than average H. mantegazzianum, might correspond to H. pubescens (Hoffm.) M. Bieb., which (according to Mandenova 1974, 1987) has shorter leaf-lobes and fewer rays than H. mantegazzianum. According to Wellendorf (1980), specimens of large Heracleum species from D were sent to I. Mandenova and identified as H. pubescens. The material that was sent to her has not been seen, but may belong to the morphotype mentioned above; however, its identity could not be confirmed, since material of H. pubescens in LE is heterogeneous, and does not correspond to Mandenova’s description.29 – Two other specimens with fewer umbel rays, from S Klm Lofta 1971 (roadside) and Upl Danderyd 1954 (railway station), deviate from both H. mantegazzianum and the alleged H. pubescens in having fruits that are almost circular in outline and have distinctly wider vittae. In Nordic material of H. mantegazzianum there is a partial correlation between leaf indumentum and the length of the hairs on rays and pedicels (specimens with glabrous leaves have shorter hairs in the inflorescence than specimens with hairy leaves). – See also under H. persicum. 27OBS ändrat igen, hoppas det blivit rätt nu då! f.d. ny text: A morphotype with no elongated leaf-lobes, umbels with fewer rays (9–40), and flowers with red-violet anthers but without distinctly radiating outer petals has been seen from D ØJy Viborg 1941 (garden relic), Odder 1971, VJy Skjern 1968 (ditch), Sjæ Gentofte 1961, Søndersø 1965 (river), Langholmen 1965, N Te Langesund 1951, Porsgrunn 2005, S Srm Nynäs 1903 and Upl Danderyd 1945, 1981(railway station). This variant, which is also somewhat smaller than average H. mantegazzianum, might correspond to H. pubescens (Hoffm.) M. Bieb. According to Wellendorf (1980), the material [om det var alla de nämnda?!] from D of large Heracelum species were claimed by I. Mandenova to be conspecific with H. pubescens, which has, among other diagnostic characters, shorter leaf-lobes and fewer rays than H. mantegazzianum (Mandenova 1974, 1987). However, herbarium material in LE is heterogeneous and does not correspond to Mandenova’s description. – There are two other specimens with fewer umbel rays, from S Klm Lofta 1971 (roadside) and Upl Danderyd 1954 (railway station), which deviate from both H. mantegazzianum and the alleged H. pubescens in having fruits that are almost circular in outline and have having distinctly wider vittae. OK, bra med smärre justeringar. OBS! Det danska materialet som Mandenova bestämt till H. pubescens har jag inte sett; jag har spekulerat över om det ev. ligger i Tbilisi där hon var verksam /LF Gammal text Several specimens (from”D ØJy Viborg 1941 (garden relic), Odder 1971, VJy Skjern 1968 (ditch), Sjæ Gentofte 1961, Søndersø 1965 (river), Langholmen 1965, N Te Langesund 1951, Porsgrunn 2005, S Srm Nynäs 1903 and Upl Danderyd 1945, 1981(railway station)”) deviate in being somewhat smaller and having leaves without elongated lobes, umbels with fewer rays (9–40), and flowers with not distinctly radiating outer petals and red-violet anthers. This morphotype might correspond to H. pubescens (Hoffm.) M. Bieb., which is characterized by having, i.a., shorter leaflobes and fewer rays than H. mantegazzianum (Mandenova 1974, 1987). However, herbarium material in LE is heterogeneous and does not correspond to this description. According to Wellendorf (1980), records from D were claimed to be conspecific with H. pubescens by I. Mandenova. Perhaps the specimens that were sent to her are conspecific with the ones mentioned above. Furthermore, two specimens from ”S Klm Lofta 1971 (roadside), and Upl Danderyd 1954 (railway station)” also have fewer rays, but deviate from both H. mantegazzianum and the formerly mentioned morphotype by having fruits that are almost circular in outline, with distinctly wider vittae. 28 struket: Vf Holmestrand 1922, - `sparad, måste kolla, kanske är det något som måste meddelas att det var fel? Ex inte helt typiskt, dessutom sannolikt cult. (laererskolen) /LF – ska föras in i geofilen OK inlagd som ‘doubtful’ i geofilen /LF 29 Förut stod det: were claimed by I.P. Mandenova to be conspecific with H. pubescens; according to her, they had more bracts and narrower leaf-lobes than H. mantegazzianum in Caucasus (Wellendorf 1980). vilket inte betyder riktigt samma sak, nya syftningen är väl inte avsiktlig? Jo, jag är osäker men undrar om inte Mandenova förväxlade karaktärer i notisen till Wellendorf ?/LF Sium 11 Similar taxa. Heracleum mantegazzianum differs in habit from H. persicum, which usually has several stems. More differences: H. persicum has a base that is ± continuously violet, more elongated leaf-blades with more (2–4) pairs of leaflets, and umbels that are usually more convex, especially at anthesis. See also the key. 3. Heracleum persicum Desf. ex Fischer F.E.L. Fischer, Ind. sem. horti Petrop. 7: 50 (1841). – Described on cultivated material (from Iran). H. laciniatum auct. scand., non Hornem. (1813). H. tromsoensis R. Elven (2005), nom. nud. D Hårfrugtet Bjørneklo. N tromsøpalme. F jättiputki. S tromsöloka. Literature. Alm & Jensen 1993, Often & Graff 1994. Hemicryptophyte (perennial). Plant to 1.8–2.8 m, with 1–5 stems; scent aromatic, reminding of anise. Stem hollow, sparsely to densely hairy with bristle-like, short hairs, sometimes with small purplish patches; basal part 30–40 mm thick, usually sulcate, usually continuously purplish, the colour gradually thinner upwards or sometimes disintegrating into small purplish spots. 30Leaves 31with a rather broad, usually purplish sheath; petiole 55–100 cm; blade 1-pinnate, 43–120 × 34–80 cm (length/width ratio 1.1–1.5), lower side densely covered with short hairs, upper side glabrous. Leaflets (1–)2–4 pairs. Apical leaflet usually 2-pinnatifid, with 2–4 pairs of primary lobes; petiolule 7–11 cm; blade 21–42 × 19–44 cm (length/width ratio 0.9–1.2); base truncate to deeply cordate; margin usually obtusely serrate; lobe apices short, broadly acute. Umbels ± convex, 10–15 cm high and 30–50 cm wide (at anthesis); 32rays straight or bent inwards (at anthesis 30 Omskrivet, kolla. OK /LF Nu undrar jag vad ”continuously” avser: ej fläckig utan enfärgat purplish, eller att den är purplish redan som ung? (troligen inte?) Jag försöker hitta annat ord för ”enfärgad”!/MA Med ‘continuosly purplish’ avser jag precis vad du skriver – att den inte är purpurprickig utan att prickarna “flyter samman” till en smet /LF (i mail 081111: ) --"continuously" används ofta om tid, därför hakar jag upp mig på denna petitess (alltså jag blir distraherad av ordet när jag läser). När det inte står något alls antar man ju normalt att det är jämnt fördelat, fast här kan man kanske tro att det är likadant purplish som högre upp. Är det en bra karaktär? "solid" används för enfärgad, men här är det nog olämpligt för man kan tro att ett komma försvunnit, och att "solid" avser att den inte är ihålig vid basen..... "evenly" påstår kanske något som inte är sant? "usually purplish all over" kanske? Karaktären är nog bra, men kan kanske specificeras lite - den kontinuerliga purpurröda färgen försvinner successivt om man följer stammen uppåt, men kan ibland upplösas i SMÅ röda prickar (syns ibland även på bladen). Hos mantegazz. är fläckarna något större och alltid tydliga. FÖRSLAG: "Stem hollow, sparsely to densely hairy with bristle-like, short hairs; basal part 30-40 mm thick, usually sulcate, usually continuously purplish, with the colour gradually disappearing or sometimes disintegrating into small purplish patches [dots?]." /LF OK - det blev rätt annorlunda, nu verkar det som om den knappast är fläckig upptill - men om inte det är missledande (jag känner inte arten!!) så är det nog bra, förutom att jag ska få in ett "upwards" eller något sånt. Kanske så: ...purplish, the colour gradually thinner upwards or sometimes disintegrating into small purplish spots. OK bra /LF081112 Patch låter för stort, dot lite för litet, spot är kanske lite mittemellan? 31 antal står på sphondylium saknar uppgift (mindre viktig karaktär) /LF 32 peduncle? Saknar uppgifter /LF Sium 12 usually distinctly inwards-bent), 8–22 cm, on the adaxial side sparsely to densely covered with triangular, whitish, ± ascending papillae, and usually also sparsely to densely covered with rigid, ± ascending hairs. Bracts 10–1833, persistent. Umbellules 35–84; pedicels 1.3–3.5 cm, with similar indumentum on the adaxial side as the rays but usually more densely hairy34. Bractlets 10–1535, persistent, 3–10 × 0.2–0.8(–1.1) mm, distinctly hairy. Flowers (34–)40–80 per umbellule; sepals 0.2–1 mm; petals 6.5–14 × 8.5–15 mm (length/width ratio 0.7–1.2), bifid (apical cut (2–)3–8 mm deep), with 3–5 mm wide lateral lobes and usually distinct veins; filaments 3–4.5 mm; anthers 0.9–1.3 mm (stamens sometimes reduced and sterile). Pollen grains 51–66 × 24–33 μm36 (length/width ratio 1.6–2.2). Fruit oblong in outline, with a dense cover of flexuose hairs longer than 1 mm, sometimes also with lateral, minute, upwards-directed hooks. Mericarps 8–14 × 6–9.5 × 0.5–1.2 mm (length/width ratio 1.2–1.8); wings 0.6–1.1 mm wide; dorsal vittae slightly expanded at the apex, the median ones (4.5–)6–8.5 × 0.4–0.9 mm, the lateral ones (4–)5.5–8 × 0.5–1 mm; stylopodium flattened, 1.7–2.8 mm wide; style 1.3–3 mm, directed ± outwards. – Early summer to mid-summer. 37 Distribution. Heracleum persicum has long been in cultivation as an ornamental in the Nordic countries (known since 1836 in N VFi Alta, probably since the 1850’s in N Tr Tromsø), but there are no records of escaped occurrences in those areas until 1901 (cf. Alm & Jensen 1993); the first verified Nordic escape record is from F U Sipoo 1871. The species is probably more common than shown in the map; only records where vouchers have been checked are included here. – D casual in Sjæ København 1987 (park), 38?NJy Koldby 1937. N resident in the north, probably scattered along the coast from VFi Hammerfest to ST Trondheim; casual in Op Lillehammer 1944 (roadside)39, He Ringsaker 1995 (roadside), Ak 40Oslo 1955, 1996 (forest fringe). S PL Arvidsjaur 1994 (Renträsk, resident at an abandoned factory); LyL Tärna 2003, Bl Sturkö since 2003 (farm). F V 5 localities, resident in Lohja since 1889 33 Obs, antal svepeblad från fältmätningar istället för herbariekollekter/LF 34 more densely than vad? than on the abaxial side? (Man undrar ett ögonblick om jämförelsen skulle vara med rays.) (den icke papillösa och mindre håriga sidan = abaxial = sidan vänd FRÅN mitten, ok?) OK, jämförelsen gäller mot flockstrålarna (justerat) /LF Nu ser jag att det inte är glasklart om hårigheten (både på rays och pedicels) är på samma sida som papillose eller runtom, och vet inte om det har någon betydelse. Jag märkte det när jag försökte ändra ordföljden - det blev olika betydelse då… papiller och hår sitter enbart på ovansidan (adaxial side) /LF Justerat förslag: with similar indumentum on the adaxial side as the rays but usually more densely hairy. /OK LF 081111 35 Obs, antal svepeblad från fältmätningar istället för herbariekollekter/LF 36 tecknet hade försvunnit Måtten stämmer /LF (Obs enhet my) /LF 37 Omskrivet. Gamla: Distribution. Heracleum persicum is known to have been in cultivation as an ornamental in the Nordic countries since 1836 in N VFi Alta, and probably also since the 1850’s in N Tr Tromsø, although no records of escaped occurrences are known in the region until 1901 (cf. Alm & Jensen 1993). The first verified Nordic record of the species as an escape is from 1871 (F U Sipoo). The species is probably more common than shown in the map; only records where vouchers have been checked are included here. – D casual in Sjæ København 1987 (park), NJy Koldby 1937?37. N resident in the north, probably scattered along the coast from VFi Hammerfest to ST Trondheim; casual in Op Lillehammer 1944 37(roadside), He Ringsaker 1995 (roadside), Ak 37Oslo 1955, 1996 (forest fringe). S PL Arvidsjaur 1994 (Renträsk, resident at an abandoned factory); casual in Bl Sturkö since 2003 (farm), LyL Tärna 2003. F V 3 localities, resident in Lohja since 1889, U 6 localities, resident in Helsinki since 1923; casual in St Vammala 1986 (courtyard), EH Tammela 1911–12 (escaped in garden), PS Heinävesi 1958 (courtyard), KP Haapavesi 1980 (naturalized), and OP Oulu 1911 (river). – Map xxx. 38 DANISH REVIEWERS: Specimen label says “Koldby”; please help with province for the locality/LF 39 struket, vet ej varför, ombestämd?: Vestre Toten 1993/MA 40 struket, vet ej varför, ombestämd?:: Frogn 1933 (spontaneous in gardens), /MA Sium 13 (cultivated in the 1880’s)41, U c. 10 localities (as a relic and in waste ground)42, resident in Helsinki since 1923; casual in St Vammala 1986 (courtyard), EH Tammela 1911–12 (escaped in garden), Parala 1956 (lakeshore), PS Heinävesi 1958 (courtyard), KP Haapavesi 1980 (naturalized), and OP Oulu 1911 (river43). – Map xxx. Iran, Iraq and Turkey. Habitat. Sun-exposed to shaded, nutrient-rich habitats. Mostly in man-made places such as roadsides, railways, ruderal places, gardens and parks; at least in N and F sometimes also in seminatural habitats such as deciduous woodland, ungrazed grassland44 and shores. At several localities, e.g. in N Tr Tromsø, forming dense and large stands. Biology. Heracleum persicum is a truly perennial herb, producing flowering shoots repeatedly for several years, and may survive long at a locality even if sexual reproduction fails (in contrast to H. mantegazzianum, which is hapaxanthic and depends on seed reproduction for survival). Pollen fertility varies extensively (see under Taxonomy)45. Seed reproduction seems to be less vigorous in H. persicum than in H. mantegazzianum, as would be expected, at least in the southern parts of the area; however, in N Tr Tromsø, seed reproduction is extensive, and H. persicum has expanded enormously (also hybridizing with H. sphondylium subsp. sibiricum). According to Often & 41 According to documentation file 1889 to 1946, is it still there? Might well be mentioned that it has obviously been grown in gardens (cultivated specimens) OK included /LF 42 Något om hur den växer? I dokumentationsfilen står ”5!”, men det verkar vara fler redan inom Helsinki. OK justerat o. inlagt “e.g. in ruderal sites and as a relic” /LF RAINO can you improve? 43 ”river” konstigt för arten – ”riverside” eller vad? Står ju Oulujoki, är det grunden för ”river”? Ja, struket /LF RAINO can you improve? 44 konstigt – meadows är hömarker, avses ”abandoned pasture”? Eller mer obestämt ”overgrowing grassland”? Ja, ‘overgrowing grassland’ ska det vara /LF ‘ungrazed grassland’ inlagt /LF ALL REVIEWERS: you are very welcome to improve… 45 OBS att preciseringarna om pollenfertilitet from 0–100% (measured as percentage of grains stained in cotton blue), on average clearly lower than in H. mantegazzianum where it is mostly above 80% nu är helt borttagna – OK eller misstag? Ja, misstag – inlagt igen /LF Detta låter för generellt, du kan bara uttala dig om vad du själv eller någon annan undersökt (baseras siffrorna förresten på egna mätningar eller borde en ref ges?) och insamlingsgraden är låg enligt tidigare avsnitt. Var finns de med nedsatt fertilitet, är det i Tromsö?? Om H. persicum exempelvis är höggradigt homozygot med dominanta drag skulle man bara sällan, i F2 och senare generationer, kunna spåra ett hybridursprung i utseendet. Räcker insamlingsgraden för att man ska kunna dra några slutsatser? Siffrorna baseras på egna mätningar, men materialet är relativt litet som du påpekar. Vissa områden har blivit rikligt företrädda i samlingarna (såsom Loja i södra Finland och Troindheim) och från dessa kan pollenfertilitet variera från 0% till över 80%. Jag tycker dock att den nedsatta pollenfertiliteten hos H. persicum jämfört med H. mantegazzianum är viktig att påpeka /LF En lite försiktigare skrivning (fortfarande kanske för tvärsäker) kunde vara ”H. persicum specimens with reduced male fertility are probably not of hybrid origin, since no correlation with any other traits have been found (examined specimens are morphologically typical). “– Detta kanske skulle stå på variation??? OK, låter rimligt (både formulering och placering) /LF OBS! Jag skulle vilja ändra formuleringen lite, dels eftersom pollenfertiliteten hos H. persicum från Turkiet är hög, och dels eftersom jag hittar några små detaljskillnader; förslag inlagt på sista stycket i ‘Taxonomy’ /LF OBS resonemanget på sphondylium om translokationsheterozygoti (vilket kanske i o f s motsäger min tanke om osynliga hybrider ) på sphondylium, finns det skäl att säga något vagt om det som en möjlig förklaring och hänvisa till sphondylium? Det är en lockande tanke, men jag tvekar – bl.a. finner jag ibland pollenkorn som sitter kvar i sina tetraedrer hos pollensterila H. persicum; jag har svårt att tro att detta skulle orsakas av translokationer enbart? /LF OBS att preciseringarna om pollenfertilitet nu är helt borttagna – OK eller misstag? Sium 14 Graff (1994), only the seeds of the primary umbel are developed in H. persicum, while those of secondary and tertiary umbels are aborted. 4647 Taxonomy.48 The Nordic material of Heracleum persicum has raised great interest and caused much confusion; it has often been taken for H. mantegazzianum. In northern N (Nordhagen 1940, Lid 1963) and in F (Hiitonen 1933) it was 46 Otydligt. Avses ”spread” eller avses frösättning? ”reproduce” bör ersättas här. Lite längre ner står om frösättning så det kanske är det som avses, men det är väl naturligt hos en äkta perenn att den har lägre frösättning än en hapaxanth och det borde skrivas som en naturlig sak. Det är väl också rätt naturligt att man får låg frösättning om pollenet är dåligt. – Om man avskriver hybridisering bör man också diskutera förklaringar till den höga fertiliteten där arterna hybridiserar. OK, justerar till “..does not seem to produce offspring as vigorously as H. mantegazzianum” /LF Förstår fortfarande inte riktigt, låter det stå kvar så länge. 47 OBS det nordnorska är väl studerat och omskrivet, referenser? Nu blev det en referens till en detalj (vilka frön som blir fullbildade), men inte generellt – ser lite mystiskt ut. Kanske något malplacé, men det funkar väl som det står? /LF Ja, eftersom den ges under Literature gör det ju det. 48 Mitt förslag var alltså det som står i själva texten, inte nedanstående – jag såg inga justeringar i själva texten, bara i fotnoten, så jag blir osäker på vilket du godkände!! ;) För säkerhets skull frågar jag om. Jag föreslår alltså följande text [magentajusteringar från LF]: The Nordic material of Heracleum persicum has raised great interest and caused much confusion; it has often been taken for H. mantegazzianum. In northern N (Nordhagen 1940, Lid 1963) and in F (Hiitonen 1933) it was earlier recognized under the name of H. laciniatum Hornem. Brummitt (1968) suggested that the Nordic material belongs to H. persicum, and that name was adopted for material from F in Hämet-Ahti (1984). Original material of H. laciniatum has not been seen by the present author, but according to the protologue in Hornemann (1813, p. 279) that species has entire, lobed leaves that are tomentose beneath, and it is not likely to be conspecific with this taxon. Øvstedal (1987) concluded that the material in northern N was not conspecific with H. persicum, after comparison with material from Turkey; he based his judgement on several differences, e.g., size and shape of the fruits, width of the vittae, shape and division of the leaves, and pollen size. Taking all the Nordic material into consideration, however, the conclusion must be that it does belong to H. persicum. However, although the Nordic taxon is somewhat variable in leaf-shape, shape and size of fruits and width of vittae, the material of H. persicum from Turkey and Iran corresponds fairly well with it in most characters. The size of the pollen grains in H. persicum (52–60 × 25–30 ×m according to Øvstedal 1987) falls within the variation range of the Nordic material. The size of the fruits and the vittae in material from Turkey and Iran are also within the Nordic variation range, although the fruits usually do not have the typical cuneate base of H. persicum from the Orient and the vittae reach proportionally further down on the mericarps (median vittae reaching downwards to 68—76% of mericarp length in Nordic material, and to 61—71% of mericarp length in oriental H. persicum). The differences in leaf shape of the Nordic material varies somewhat but corresponds fairly well with may be explained by the fact that the leaves from the Nordic material in Øvstedal (1987) derived from a higher position on the stem than those of the studied specimens of H. persicum from Turkey and Iran. Furthermore, important diagnostic characters not included in Øvstedal (1987), such as the shape of the dorsal vittae and the indumentum on rays and fruits, also agree with the oriental material from Turkey. 48 It cannot be excluded that the Nordic taxon is of hybrid origin. It could thus have arisen by hybridization between H. persicum and H. mantegazzianum. The variation is extensive in the Nordic taxon, with no clear limit towards H. mantegazzianum, and it also shows a high degree of reduced pollen fertility also in fairly typical specimens (material from Turley has a high pollen fertility). However, H. mantegazzianum was not in cultivation in Europe until the end of the 19th century, why at least older records should not have been influenced by this species. Thus, it is more plausible that the Nordic material derive from H. persicum of SW Asia, and may have acquired the differences from this species, e,g,, by genetic drift. inte nedanstående. /MA gamla texten: Taxonomy. The identity of this taxon has hitherto been uncertain. It has largely been confused with H. mantegazzianum, and was earlier recognized under the name of H. laciniatum Hornem. in northern N (Nordhagen 1940, Lid 1963) and in F (Hiitonen 1933). Although original material of H. laciniatum has not been seen, according to the protologue in Horneman (1813, p. 279) the species has entire, lobed leaves that are tomentose beneath, and it is not likely to be conspecific with this taxon. Sium 15 earlier recognized under the name of H. laciniatum Hornem. Brummitt (1968) suggested that the Nordic material belongs to H. persicum, and that name was adopted for material from F in Hämet-Ahti (1984). 49 Original material of H. laciniatum has not been found by the present author, but according to the protologue in Hornemann (1813, p. 279) that species has entire, lobed leaves that are tomentose beneath, and it is not likely to be conspecific with this taxon. 50Øvstedal (1987) concluded that the material in northern N was not conspecific with H. persicum; he based his judgement on comparison with H. persicum material from Turkey differing in several characters, e.g., size and shape of the fruits, width of the vittae, shape and division of the leaves, and pollen size. When more of the Nordic material is taken into account, and also H. persicum material from Iran51, a comparison will show fairly good correspondence in most characters. Nordic fruits usually do not have the typical cuneate base of H. persicum from the Orient, and their vittae are proportionally slightly longer (the length of vittae in oriental H. persicum is 61–71% of the mericarp length, in Nordic material 68–76%). However, in leaf shape, in absolute size of fruits and vittae and in size of the pollen grains (52–60 × 25–30 μm in H. persicum according to Øvstedal 1987), the restricted Turkish and Iranian H. persicum material studied falls within the greater variation range of the Nordic material. Also in important diagnostic characters not mentioned by Øvstedal, such as the shape of the dorsal vittae and the indumentum on rays and fruits, the Nordic taxon agrees with oriental H. persicum material. Nordic H. persicum varies extensively, with no clear limit towards H. mantegazzianum. Even in fairly typical specimens the pollen fertility is frequently reduced (0–100%; above 80% in H. persicum from Turkey and in H. mantegazzianum), which suggests a hybrid origin. H. persicum is here defined as having stems, petioles and rachises In Brummitt (1968), H. persicum was suggested to be conspecific with the Nordic material. Øvstedal (1987) compared material from northern N with H. persicum from Turkey. He found several differences, e.g., size and shape of the fruits, width of the vittae, shape and division of the leaves, and pollen size, and he concluded that the material of northern N was not conspecific with H. persicum. The Turkish material has been reinvestigated, taking into consideration However, when taking all Nordic material of the studied taxon into consideration, The size of the fruits and the vittae in H. persicum from Turkey falls within the variation range of it the Nordic material. The differences in leaf shape could perhaps be explained by the fact that the leaves from the Nordic material in Øvstedal (1987) derived from a higher position on the stem than the leaves from specimens of H. persicum. The size of the pollen grains in H. persicum (52–60 25–30 m) also falls within the variation range of the Nordic material. Furthermore, important diagnostic characters of the Nordic taxon, such as the type of indumentum on the rays and the fruits, and the shape of the dorsal vittae (not included in Øvstedal 1987), are also in correspondence with the material from Turkey. It can thus be concluded that the Nordic taxon is conspecific with H. persicum. OK med smärre justeringar /LF 49 I gamla texten fanns följande: A picture of a herbarium specimen of H. persicum in Mandenova (1987) corresponds very well with the Nordic material, both in leaf and fruit characters. Har du senare sett material som är “tyngre”? Jag kan inte utläsa det av texten, därför undrar jag. Ja, det är riktigt – bilden gav mig först en indikation som kunde bekräftas när jag sett material /LF 50 Några kommentarer bevarade från arbetet med det här avsnittet: The variation is extensive in the Nordic taxon, with no clear limit towards H. mantegazzianum, and it also shows a high degree of reduced pollen fertility also in fairly typical specimens (material from Turley has a high pollen fertility). It cannot be excluded that the Nordic taxon originated from hybridization between H. persicum and H. mantegazzianum. However, H. mantegazzianum was not in cultivation in Europe until the end of the 19th century, why at least older records should not have been influenced by this species. Thus, it is more plausible that the Nordic material derive from H. persicum of SW Asia, and may have acquired the differences from this species, e,g, by genetic drift. Tveksam slutsats, om allt annat pekar mot hybridogent ursprung/introgression, vilket jag tycker det verkar göra. Vår historiska dokumentation är så bristfällig! Det kan ju ha varit hybrider man först fick in???? Till exempel. Tetx justerad (se ovan) /LF jag har bl.a. fyllt ut med mantegazzianums senare introduktion, samt ytterligare möjligheter till variationen: hybridisering med andra arter, och även i ursprungsländerna (OBS! dock att H, persicum inte är känd i Kaukasus, där H. mantegazzianum är endemisk). Förhoppningsvis en förbättring, men jag inser att det haltar lite – bl.a. förklarar ‘genetic drift’ inte den reducerade pollenfertiliteten, men det gör kanske inte så mycket. /LF …… det är svårt att dra gränsen mellan variation <inom> H. persicum s.str. och variation mot intermediärer med H. mantegazzianum, så jag vill gärna inte gå in för mycket i detalj med att ange VAD det är som varierar. Sålunda vill jag ta bort [(especially in leaf-shape, shape and size of fruits and width of vittae),]. För övrigt OK /Lars 091112 i mail 51 Har du sett på de turkiska arken Övstedal studerade? Jag har lånat just det material från Turkiet som Øvstedal tidigare studerat /LF Sium 16 without large red spots, leaves with crenate margins, rays and pedicels with whitish, acute, antrorse papillae and densely long-hairy ovaries/fruits. Specimens deviating in any of these characters have been classified as intermediates and treated as H. mantegazzianum × persicum. However, the true nature of the intermediates is uncertain. H. mantegazzianum is a Caucasian endemic (H. persicum is not known from Caucasus) which was discovered and brought to European gardens in the 1890’s. It was first collected in Norden in 1896 (in S Srm; it was not collected until after 1950 in the other Nordic countries). Thus, the older Nordic material would not be expected to include any intermediates, but it does (e.g., from D Sjæ 1869 and 1888). Other possible explanations to the great variation in Nordic H. persicum may be hybridization with other allied species, either at their original sites or in cultivation. Also, genetic drift may have added to the variation seen today. Variation. A deviating morphotype has been collected in F (EH Janakalla 1959, Luopionen 1966) and S (Nb Edefors 1997 and LyL Lycksele 1993, both in grassland). It has more densely hairy lower leaf surfaces; the rays and pedicels lack hairs and have papillae that are similar to those in H. mantegazzianum, and the ovaries/fruits have shorter hairs than H. persicum. It might correspond to the Russian species H. sosnowskyi Mand. (cf. Leht 1999, Mandenova 1974), but does not agree with material seen of H. sosnowskyi from Estonia. – See also Variation under H. mantegazzianum.52 Similar taxa see Heracleum mantegazzianum (2). 4. Heracleum platytaenium Boiss. Boissier, Ann. Sci. Nat. ser. 3 (1): 331 (1844). – Described from Turkey Literature. Davis 1972. Hemicryptophyte (short-lived hapaxanth). Plant up to 2 m; scent aromatic. Stem hairy, with bristle-like hairs, hollow; basal part at least 20 mm thick, sulcate. Leaves with a rather short sheath; petiole 15–30 cm; blade large, simple or 1pinnate with 1 pair of leaflets53, lower side with a dense cover of weblike hairs, upper side sparsely bristly. Apical leaflet 2-pinnatifid, with 2–3 pairs of primary lobes; petiolule 5–10 cm; blade 21–24 × 24–25 cm (length/width ratio 0.8–1); base cordate; lobe apices rounded; margin dentate with small teeth. iv Umbels convex, 10 cm high and 25 cm wide. Bracts 0 . Umbellules 34–42; rays straight, 11.5–16 cm, on the adaxial side with ± antrorse54, hyaline or whitish papillae55, without hairs. Bractlets 12, rather short. Flowers 48–60 per umbellule; pedicels 1.7–2.2 cm, with smaller and more obtuse papillae on the adaxial side; petals 2.5–5.5 × 1.5–3.5 mm, bifid (apical cut 1.2–3 mm deep); filaments 4–4.5 mm; anthers 0.9–1.2 mm. Pollen grains 65–67 × 30–34 μm. Fruit oblong in outline, with a sparse to dense cover of c. 0.5 mm long glandular hairs. Mericarps 9.5–13.5 × 7.5–10.5 × 0.8–1 mm (length/width ratio 1.3–1.4); wings 0.9–1.2 mm wide; dorsal vittae broad, the median ones 6.5–7 × 1.2–1.5 mm, the lateral ones 5.5–6 × 1.2–1.5 mm; stylopodium low-conical, 1.7–2 mm wide; style 3–3.5 mm, deflexed. – Early summer. 52 MAGDALENA: stycket omskrivet. – Luopionen 1966 en referens? Lägg till i littlistan. Hänvisningen till mantegazzianum förstår jag inte, där står väl inget ytterligare (den intermediära behandlas ju här)? Nej, Luopionen är en LOKAL! (text justerad) - Hänvisningen till H. mantegazzianum beror på att där beskrivs ytterligare avvikande morfotyper, vilka kan vara relevanta att läsa om även i anknytning till H. persicum (jag ger ett förtyligande) /LF 53 ej jämförbart med de andra arterna Material ytterst bristfälligt; jag ger enbart en generell beskrivning av storleken /LF 54 ascending låter lite skumt om så små ting som papiller! Är det rätt ord? kanske ‘antrorse’? /LF 55”translucent” betyder ju bara ”som man kan se ljus igenom”, men det kan ha olika färger inkl. vitt. Så detta blir lite förbryllande! Avses ”vattenklara” (färglösa kontra vitaktiga) (då ska det nog stå ”hyaline”)? on the adaxial side with ± antrorse, hyaline to whitish papillae, without hairs. OK, det blir bra /LF Eller ”genomskinliga” (translucent kontra opaka) (då kanske ”whitish” räcker? Eller är det diagnostiskt att de kan vara allt från translucent till opaka)? on the adaxial side with ± antrorse, whitish papillae, without hairs. Sium 17 Distribution and habitat. Grown as an ornamental; probably escaped or remaining from cultivation. – D Sjæ Jonstrup 1886, Roskilde 1950 (road embankment), København 1981. S Gtl Hörsne 2004 (large population on roadside since 1950’s), Nrk Askersund 1990–200156 (garden relic). – Map xxx. Turkey. Hybrids 57Heracleum mantegazzianum × persicum. – Specimens intermediate between H. mantegazzianum and H. persicum are referred to this hybrid, although the hybrid origin is uncertain (see above under ‘the giant species’). Variable, and sometimes different from both alleged parents: the leaves are frequently more densely hairy, and the umbels have sometimes fewer rays (less than 35). Differences from H. mantegazzianum: red spots on stem, petioles and rachises sometimes lacking, leaves have usually no elongated lobes, and rays and pedicels have ± acute papillae. Differences from H. persicum: stems, petioles and rachises are sometimes redspotted, leaf-margins are more finely toothed, papillae on rays and pedicels are weaker, and fruits have shorter hairs. Specimens seen from most of continental Norden, but apparently very scarce. Resident in D Sjæ 6 localities (e.g. Roskilde, large population on seashore; reported as H. persicum58). N Ak 4 localities, He Eidskog and Os, ST 10 localities, NNo Beiarn and Bodø, Tr Tromsø, Balsfjord and Lyngen. S Upl Stockholm (Kungsholmen; reported by Lindberg 1983 as H. stevenii). F V Lohja, U Helsinki (Tuomarinkylä). Heracleum persicum × sphondylium. – N polarpalme. – Variable, with some specimens more similar to H. persicum and others similar to H. sphondylium. Leaf-blades intermediate in stiffness, upper side glabrous or with very few, minute bristles, especially along the edge, lower side with rather sparse hairs. Rays and pedicels covered with opaque, acute, patent to ascending papillae and 56 två olika förekomster alltså? Två olika insamlingar, men sannolikt från samma bestånd /LF FRÅGA TILL LARS LÖFGREN BÖR KUNNA UNDANRÖJA DETTA/MA 2008 Mailsvar från Lars Löfgren: “tycks sedd på Rymans lokal” – bör sålunda vara samma bestånd, varför jag ger intervall /LF 57 Jag förväntar mig en hybridrubrik ”mantegazzianum x persicum”! OK inlagt /LF Du har ju en stor mängd ”intermediates” mellan mantegazzianum och persicum. Eftersom dessa redovisas som separata arter finns det väl anledning att redovisa det intermediära materialet – och att diskutera vad det är för något. Sedan är det OK om det står att det intermediära kanske inte alls är hybrider etc, men hur man skiljer ut arterna från det du betraktat som intermediärer bör stå och det verkar mest praktiskt att ha det här. Jag vill se Similar to…. but…. eller Differs from… in… eller Differences from H. persicum: …. Differences from H. mantegazzianum: …. Jfr t.ex. på Salix! Det är nu lite oklart vad som avses åtminstone under bladen (är det elongated lobes eller without elongated lobes som är som H. mantegazzianum?, det måste jag kolla) Mantegazz har ‘elongated lobes’, men intermediärerna har normalt inte det /LF Antalet rays: ”många” verkar mer mantegazzianum, är det korrekt uppfattat att du tänkte så? Båda arterna brukar oftast ha mycket mer än 35 så det är rätt vagt… Nej, ‘mostly with many rays’ innebär att de mestadels har många flockstrålar som mantegazz och persicum, MEN kan ha färre än vad dessa har, dvs. färre än 35 /LF Följande är bara ett första försök – gulfärgade karaktärer har jag inte jämfört med arterna eller vet inte hur jag ska hantera. Heracleum mantegazzianum persicum. – Specimens intermediate between H. mantegazzianum and H. persicum are referred to this hybrid, although they may not be of hybrid origin (see above under ‘the giant species’). – Variable; with or without red spots on stem, petioles and rachises; leaves frequently fairly densely hairy beneath. Similar to H. persicum, but the leaves have more finely toothed margins, and no elongated lobes, the rays and pedicels have weaker, ± acute papillae, and the fruits often have hairs of the same length as in H. mantegazzianum Umbels mostly with many rays (but sometimes fewer than 35). Om jag spaltar upp i karaktärsskillnader från ‘föräldrarna’ , skulle det kunna bli så här: Differs from H. mantegazzianum by sometimes lacking red spots on stem petioles and rachises, leaves usually without elongated lobes, and rays and pedicels with ± acute papillae. Differs from H. persicum by sometimes having red spots on stems, petioles and rachises, more finely toothed leaf-margins, weaker papillae on rays and pedicels, and fruits with shorter hairs. The leaves are frequently more densely hairy than in both ‘parents’ and the number of rays in the umbels is variable (sometimes fewer than in the ‘parents’). 58 menar du alla danska lokaler eller just Roskilde-strandpopulationen? Om det sistnämnda föreslår jag ”(….seashore, reported as…)” eller ….published as. Samma på Kungsholmen. Ref? Justerat så /LF Sium 18 sometimes also rather rigid, opaque hairs. Fruit usually with long hairs and short papillae mixed on the surface; mericarps 9–12 × 7– 9 mm; median vittae 0.2–0.4 mm wide; lateral vittae 0.3–0.7 mm wide. N ST Trondheim since 1975 (5 localities), Tr Tromsø since 1956 (resident and increasing, = H. persicum × sphondylium subsp. sibiricum)59. S Bl Sturkö since 2003 (= H. persicum × sphondylium subsp. sibiricum)60. F St Vammala 1986. Heracleum mantegazzianum × sphondylium. – Similar to H. persicum × sphondylium, but stem and petioles ± red-spotted. Rays and pedicels densely covered with translucent, patent papillae and soft, flexuous glandular hairs (no opaque, acute papillae). S Sk Lund 2001 (spontaneous in botanical garden; = H. mantegazzianum × sphondylium subsp. sphondylium). – Records from N ST Trondheim were redetermined to H. persicum × sphondylium. 59 LARS: Är den fertil? Eller vegetativt spridd????? Hybridbestånden i Tromsø är säkerligen av hög fertilitet; de är vanliga och varierar med alla övergångsformer och karaktärskombinationer från föräldraarterna. Torbjörn Alm är bekymrad över att även hybriden kan bli invasiv i Tromsø i framtiden. Vegetativ förökning känner jag dock inte till inom släktet. /LF 60 Varför två årtal – väl skilda lokaler? Är båda med subsp sibiricum? Hybriderna är kanske inte samma individ men beståndet är detsamma, med föräldraarterna bredvid (för H. persicums del är det definitivt samma individ som står kvar) /LF ger intervall istället /LF Är det borta nu? ”known since 2003” bättre om det finns kvar Nej, finns sannolikt kvar; text justerad /LF Crithmum 19 References References Heracleum Alm, T. & Jensen, C. 1993: Tromsø palmen (Heracleum laciniatum auct. scand.) – noen kommentarer til artens innkomst og ekspansjon i Nord-Norge. Blyttia 51: 61–69. Brummitt, R.K. 1968: Heracleum L. In T.G. Tutin et al. (eds), Flora Europaea 2: 364–366. Cambridge. Davis, P.H. 1972: Heracleum L. In P.H. Davis (ed.), Flora of Turkey 4: 488–500. Edinburgh. Hämet-Ahti et al. 1998: Retkeilykasvio, ed. 2. Helsinki. Hansen, S.O., Hattendorf, J., Nielsen, C., Wittenberg, R. & Nentwig, W. 2007: Herbivorous arthropods on Heracleum mantegazzianum in its native and invaded distribution range. In Pysek, P., Cock, M.J.W. & Ravn, H.P. (eds.), Ecology & Management of Giant Hogweed: 170–188. Hattendorf, J., Hansen, S.O. & Nentwig, W. 2007: Defence systems of Heracleum mantegazzianum. In Pysek, P., Cock, M.J.W. & Ravn, H.P. (eds.), Ecology & Management of Giant Hogweed: 209–225. Hiitonen, I. 1933: Suomen Kasvio. Helsinki. Hornemann, J.W. 1813: Tyronum et Botanophilorum. København. Leht, M. (ed.) 1999: Eesti taimede maaraja [Field-guide of Estonian plants]. Estonian Agricultural University. Tartu. Lid, J. 1963: Norsk og svensk flora. Ed. 1. Oslo. Lid, J. & Lid, D.T. 2005: Norsk flora. Ed. 7 revised by R. Elven. Det norske samlaget, Oslo. Lindberg, P.S. 1983: Stockholmsfloran. Stockholm. Lundström, H. 1989: Jättelokan i Skåne. Skånes Natur, Årsbok 1989: 132–138. Madsen, H.E. Svart & Lyck, G. 1991: Introducerede planter – forvildede og adventive arter. Inst. for økologisk botanik, Københavns universitet & Skov- og Naturstyrelsen, Hørsholm. [Stencil.] Mandenova, I. 1974: Heracleum L. In B.K. Shishkin (ed.), Flora of the U.S.S:R. 17: 223–259. Mandenova, I. 1987: Heracleum. In K.H. Rechinger (ed.), Flora Iranica 162: 492–502. Moravcová, L., Pysek, P., Krinke, L., Pergl, J., Perglová, I. & Thompson, K. 2007: Seed germination ,dispersal and seed bank in Heracleum mantegazzianum. In Pysek, P., Cock, M.J.W. & Ravn, H.P. (eds.), Ecology & Management of Giant Hogweed: 74–91. Nordhagen, R. 1940: Norsk flora – Med kort omtale av innførte treslag, pryd- og nytteplanter. Oslo. Often, A. & Graff, G. 1994: Skillekarakterer for kjempebjørnekjeks – Heracleum mantegazzianum – og tromsøpalme – H. ‘laciniatum’. Blyttia 52: 129–133. Øvstedal, D.O. 1987: Er tromsøpalma sit namn Heracleum persicum Desf. Polarflokken 11: 25–26. Pysek, P., Cock, M.J.W. & Ravn, H.P. 2007: Ecology & Management of Giant Hogweed. Oxfordshire. Seier, M. & Evans, H.C. 2007: Fungal pathogens associated with Heracleum mantegazzianum in its native and invaded distribution range. In Pysek, P., Cock, M.J.W. & Ravn, H.P. (eds.), Ecology & Management of Giant Hogweed: 189–208. Tiley, G.E.D., Dodd, F.S. & Wade, P.M. 1996: Biological flora of the British Isles: Heracleum mantegazzianum Somm. & Lev. Journal of Ecology 84: 297–319. Weimarck, G. 1978: B chromosomes in Heracleum sphondylium s. lat. in Europe. In A.M. Cauwet-Marc & J. Carbonnier (eds), Les Ombellifères; contributions pluridisciplinaire à la systematique: 293–303. Perpignan. Weimarck, G., Stewart, F. & Grace, J. 1979: Morphometric and chromatographic variation and male meiosis in the hybrid H. mantegazzianum × sphondylium (Apiaceae) and its parents. Hereditas 91: 117–127. Wellendorf, M. 1980: Bjørneklo. URT 4: 89–91. i …gruppen i sin helhet introducerades under tidigt 1800-tal; bl.a. tror jag att H. persicum och H. platytaenium figurerade då (Hornemanns material av "H. giganteum" från Københavns bot. trädgård motsvarar faktiskt H. platytaenium). H. mantegazzianum beskrevs 1895 på material i en bot. trädgård i norra Italien (den fick namnet efter trädgårdsmästaren Mantegazzi, som på försättsbladet till publikationen figurerar med en jätteloka bredvid sig). Jag kan citera vad Dmitry Geltman skriver i 'Giant hogweed-volymen' om artens upptäckt: "It is relevant to point out that the majority of the pre-1872 Caucasian collections come from the Transcaucasia and Central Greater Caucasus. The Western Greater Caucasus was closed to expeditions because of war. It is for Crithmum 20 this reason that large hogweeds collected at that time mainly belong to H. sosnowskyi (in terms of the modern nomenclature), because H. mantegazzianum Somm. et Levier s.str., occurring in Western Greater Caucasus, was not then known to science. In addition, expeditions to high mountain areas were ardous and expensive. Therefore, it is not surprising that H. mantegazzianum was discovered much later, by the Caucasian expedition of S. Sommier and E. Levier in 1890 (Sommier & Levier 1895, 1900)." /LF 081112 ii Kommentar att observera vid senare redigering: [står det nåt om det eller nåt annat om Mandenova-material nånannanstans? Hänvisning här isåfall! Ev kan detaljer flyttas dit, i kommentarerna här finns en del som jag finner intressant] iii tips och crenate går dåligt ihop för mig. /MA ‘tips’ avser den förhårdade, vita/violetta spetsen av varje krenulerad tand /LF På bild av ssp sphondylium ser det verkligen ut som ”crenate”, men det ser så konstigt ut med ”tips”! OLÖST! iv THOMAS: Är detta en störande inkonsekvens mot andra familjer?