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Sendtnera 5: 35-241 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at 35 Die infragenerische Gliederung der Gattung Heliotropium L. und ihre Stellung innerhalb der subfam. Heliotropioideae (Schrad.) Arn. (Boraginaceae) H. FÖRTHER Abstract: FÖRTHER, H.: Die infragenerische Gliederung der Gattung Heliotropium L. und ihre Stellung innerhalb der subfam. Heliotropioideae (Schrad.) Am. (Boraginaceae). - Sendtnera 5: 35-241. 1998. ISSN 0944-0178. Based on extensive morphological and anatomical investigations a new classifi- cation of the Boraginaceae subfamily Heliotropioideae is given. It now includes the 8 genera Argusia Böhm., Ceballosia Kunkel, Heliotropium L., Hilgeria Förther [gen. nov.], Ixorhea Fenzl, Nogalia Verde, Schleidenia Endl. [reinstalled] and Tournefortia L. Detailed descriptions of the Boraginaceae, the subfamily Helio- tropioideae and all genera belonging to the latter are provided, supplemented by a concise determination key to all genera within the subfamily Heliotropioideae and sections within Heliotropium. The genus Heliotropium is segregated into 19 sections (sect. Chamaetropium, sect. Cochranea, sect. Coeloma, sect. Heliotham- nus, sect. Heliotrophytum, sect. Heliotropium, sect. Hypsogenia, sect. Moniman- tha, sect. Odontotropium, sect. Orthostachys, sect. Plagiomeris, sect. Platygyne, sect. Pleurolasia, sect. Pseudocoeloma [sect, nov.], sect. Pterotropium, sect. Rutidotheka, sect. Schobera, sect. Tiaridium, sect. Zeylanica [nom. nov.]). The characters used for the infrageneric classification of Heliotropium are discussed. Every section is described in detail, synonymy of the sectional names and the species belonging to them are listed, the differential characters and the relation- ships to other sections are discussed. Supplementary in the appendix all taxa ever described within Heliotropium worldwide or described within other genera, but really belonging to Heliotropium, are compiled in a comprehensive list including the complete references, types and currently accepted names. More than 150 taxa are lectotypified the first time. Zusammenfassung: Basierend auf umfangreichen morphologischen und anatomischen Untersuchungen wird eine neue Einteilung der Boraginaceae subfam. Heliotropioideae vorgestellt, nun aus 8 Gattungen bestehend {Argusia Böhm., Ceballosia Kunkel, Heliotropium L., Hilgeria Förther [gen. nov.], Ixorhea Fenzl, Nogalia Verde, Schleidenia Endl. [erneut abgetrennt] and Tournefortia L.). Die Familie der Boraginaceae, die Unter- familie der Heliotropioideae und die zur letzteren gehörenden Gattungen werden ausführlich beschrieben. Ein Bestimmungsschlüssel ermöglicht es, die Gattungen der Unterfamilie der Heliotropioideae und die Sektionen der Gattung Heliotropium anzusprechen. Für die Gattung Heliotropium wird eine Unterteilung in 19 Sektionen vorgeschlagen, bestehend aus sect. Chamaetropium, sect. Cochranea, sect. Coelo- ma, sect. Heliothamnus, sect. Heliotrophytum, sect. Heliotropium, sect. Hypso- genia, sect. Monimantha, sect. Odontotropium, sect. Orthostachys, sect. Plagio- meris, sect. Platygyne, sect. Pleurolasia, sect. Pseudocoeloma [sect, nov.], sect. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at 36 Pterotropium, sect. Rutidotheka, sect. Schobera, sect. Tiaridium und sect. Zeyla- nica [nom. nov.]. Die für die Sektionseinteilung herangezogenen Merkmale werden diskutiert. Jede Sektion wird ausführlich beschrieben, die Synonyme jedes Sektions- namens werden angegeben, eine Anzahl zugehöriger Arten wird gelistet und die Unterscheidungsmerkmale und die Verwandtschaft zu den übrigen Sektionen dis- kutiert. Im Anhang werden alle Taxa aufgeführt, die unter Heliotropium oder einer anderen Gattung beschrieben wurden; angegeben wird dabei die korrekte Autoren- schaft, das Datum und der Ort der Veröffentlichung, alle zugehörigen Typus- elemente, die Herbarien und im Falle von Synonymen der korrekte Name, unter dem das Taxon anerkannt wird. Für mehr als 150 Taxa wird zum ersten Mal ein Lectotypus angegeben. Inhaltliche Gliederung 1 Einleitung 38 2. Material und Methoden 38 3. Die Familie der Boraginaceae 40 3.1. Systematische Stellung im Pflanzenreich 40 3.2. Allgemeine Familienmerkmale 42 3.3. Gliederung der Boraginaceae 42 3.4. Nomenklatur der Boraginaceen-Unterfamilien 43 3.5. Schlüssel der Boraginaceen-Unterfamilien 44 4. Die Unterfamilie der Heliotropioideae 45 4. 1 . Merkmale der subfam. Heliotropioideae 45 4.2. Gliederung der subfam. Heliotropioideae 45 4.3. Gattungsschlüssel der Boraginaceae subfam. Heliotropioideae mit besonderer Berücksichtigung der neuen Sektionseinteilung der Gattung Heliotropium 46 5. Die Gattung Heliotropium 48 5.1. Etymologie, Symbolik, Nutzen 48 5.2. Gattungsdiagnose 51 5.2. 1 . Verbreitung der Gattung Heliotropium 54 5.2.2. Allgemeinbiologische Daten 55 5.3. Merkmalsbewertung für die Sektionseinteilung 57 5.3.1. Lebens- und Wuchsform 58 5.3.2. Sprossaufbau und Verzweigung 59 5.3.3. Indument 60 5.3.4. Blätter 61 5.3.5. Infloreszenz und Infloreszenzaufbau 61 5.3.6. Blütenmerkmale (Kelch, Krone, Griffel-ZNarbenkomplex, Staubblätter und Pollen) . 62 5.3.7. Fruchtmerkmale 64 . © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at 37 5.4. Die Sektionseinteilimg der Gattung Heliotropium 65 5.4.1. Geschichte der infragenerischen Gliederung 65 5.4.2. Graphische Darstellung der neuen Sektionseinteilung 68 5.4.3. Die Sektionen der Gattung Heliotropium 70 - Chamaetropium 70 - Cochranea 72 - Coeloma 74 - Heliothamnus 77 - Heliotrophytum 80 - Heliotropium 82 - Hypsogenia 88 - Monimantha 90 - Odontotropium 92 - Orthostachys (subsect. Bracteata & subsect. Ebracteatä) 94 - Plagiomeris 102 - Platygyne 1 03 - Pleurolasia 106 - Pseudocoeloma 110 - Pterotropium 113 - Rutidotheca 116 - Schobera 119 - Tiaridium 120 - Zeylanica 123 6. Weitere Gattungen der subfam. Heliotropioideae 126 6. 1 Argusia 1 27 6.2. Ixorhea 127 6.3. Nogalia 127 6.4. Tournefortia 128 6.5. Ceballosia 128 6.6. Die neue Gattung Hilgeria 131 6.7. Die Wiedereinführung der Gattung Schleidenia 133 7. Zusammenfassung 136 8. Literaturverzeichnis 139 9. Anhang 177 9.1. Liste aller unter Heliotropium valid publizierter Taxa 177 9.2. Liste aller unter anderen Gattungen publizierter aber zu Heliotropium gehöriger Taxa (incl. der nächstverwandten Gattungen Ceballosia, Hilgeria [gen. nov.], Nogalia und Schleidenia) 228 9.3. Nomina nuda bzw. Fehlinterpretationen, die Heliotropium-Taxa. bezeichnen 238 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.biologiezentrum.at 38 1. Einleitung Innerhalb der Pflanzenfamilie der Boraginaceae, zu deutsch der Rauhblatt- oder Borretsch- gewächse, stellt die Gattung Heliotropium L. mit ihren etwa 300 Arten die zweitgrößte Gat- tung dar. Obwohl die Zugehörigkeit zu den Boraginaceen anhand charakteristischer Familien- merkmale relativ leicht kenntlich ist, resultiert doch innerhalb der Gattung aus einer gewissen Armut an aussagekräftigen Differentialmerkmalen eine fast unglaubliche Vielfalt an taxonomi- schen Problemen. Aufgrund des enormen Gattungsumfanges und der weltweiten Verbreitung gibt es keine brauchbare, moderne Gesamtrevision. Als Vorbereitung einer derartigen Mono- graphie wünscht man sich daher für eine solche Gattung wenigstens eine Unterteilung in überschaubare Gruppen oder Einheiten. Daher die Frage: Soll man nun die beinahe unüber- sichtlich große Gattung Heliotropium nur der besseren Übersichlichkeit wegen mit einer infra- generischen Gliederung unterschiedlicher Rangstufen versehen oder, wie immer wieder gesche- hen, in kleinere Gattungen aufspalten? Bei der Beantwortung dieser Frage findet man sich schnell in einem subjektiven Zugzwang. Ein wünschenswertes Ziel heutiger Taxonomie ist es, ein natürliches und kein künstliches System zu schaffen, d.h. auch auf Gattungsebene die rea- len Verwandtschaftsverhältnisse als Grundlage jeglicher Einteilung zu verwenden. Doch bei- derart vielen Vertretern einer Gattung ist es beinahe unmöglich, umfassend repräsentative molekular-biologische, chemotaxonomische, karyologische oder sonstige aussagekräftige Un- tersuchungen durchzuführen, um eine natürliche Verwandtschaft zu ergründen. Weltweit befindet sich nahezu kaum eine Handvoll Arten in Kultur; es fehlen also jegliche Grundlagen, derartig verwertbare Daten zu erhalten. Da ein System aber in erster Linie einem gewissen Pragmatismus dient, soll hiermit erstmals der Versuch unternommen werden, die Gattung Heliotropium basierend auf klassischen, morphologisch-anatomischen Untersuchungen korre- liert mit Verbreitungsmustern ganzheitlich und übersichtlich zu gliedern und die Gesamt- gattung im Vorgriff auf künftige Teilrevisionen so umfassend natürlich gegliedert darzustellen als nach derzeitigem Kenntnisstand möglich. Mit Ausnahme der Arktis vmrden von den verschiedenen Heliotropium-Arten alle Kon- tinente erobert. Die heutigen Verbreitungsschwerpunkte liegen in den tropischen und warm- temperierten Zonen von Afrika, Vorderasien, Australien und Amerika. Von den Meeresküsten bis in höchste Gebirgsregionen, von den Wüsten und Halbwüsten bis in die feuchten Tropen sind nahezu alle terrestrischen Habitate besiedelt worden. Neben wenigen Kosmopoliten sind zahlreiche extreme Lokalendemiten zu verzeichnen. Größenverhältnissse, Wuchsform und Habitus fungieren als wichtige Parameter für ein Verständnis der Aut- und Synökologie sowie der herrschenden Selektionsdrücke. So reicht beispielsweise beim Wuchsformspektrum die Variabilität von wenigen Zentimeter großen Annuellen
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