A bibliographic database for the genus Lathyrus compiled by Dirk Enneking Co-operative Research Centre for Legumes in Mediterranean Agriculture, 1998 Occasional publication No 18 International Center for Agricultural Research in the Dry Areas, Aleppo, Syria ISSN 1-320-366 ISBN 0-86422-829-5 i Toxicity The genus Lathyrus The presence of toxic non protein amino acids (NPAAs) in the seeds of Lathyrus species have restricted their agricultural development in several countries In response to an ever increasing global demand for food and feed resources and the need to diversify modern cropping systems, the legume genus Lathyrus is receiving increased attention by agricultural scientists. Kupicha (1976) lists 152 species in the genus (Table 2) which is distributed from the sub-arctic to the subtropics. It includes a range of grain, forage, pasture and ornamental crops (Table 1). The best known species include the ornamental sweet pea (Lathyrus odoratus) and the food legume grasspea (Lathyrus sativus). Chemotaxonomic studies in the early 1960s established the presence of several toxic amino acids in the seeds of different taxonomic groups. This work provided a useful frame of reference to delineate groups of species with different seed toxins. Three NPAAs of concern are the neurotoxins Betaoxalyl-diamino-propionic acid (Beta-ODAP) (L. sativus), Diamino-butyric acid (DABA) (L. sylvestris) and the nitrile containing beta-amino-propionitrile (BAPN) (L. odoratus). Table 1 Cultivated Lathyrus species (adapted from Kearney & Smartt, 1995) Species L. annuus L. aphaca L. blepharicarpus L. cicera L. clymenum L. gorgoni L. hirsutus L. latifolius L. ochrus L. odoratus L. pratensis L. rotundifolius L. sativus L. sylvestris L. tingitanus L. tuberosus Use F, P F P F, P F, P F F O F, P O F O F, P F F T The bone deforming (osteolathyrogenic) properties of Lathyrus odoratus are due to the presence of BAPN. This compound affects the cross-linking of collagen during bone and connective tissue formation. The resultant disease is known as osteolathyrism. Recent studies in Bangladesh suggest that a metabolic precursor for this compound, 2-cyanoethyl-isoxazolin-5-one is present in the vegetative parts and immature seeds of L. sativus. It appears responsible for the osteolathyrogenic symptoms observed in some neurolathyrism patients who had consumed vegetative parts of L. sativus. The neurotoxicity of Lathyrus sylvestris and related species is caused by the toxic NPAA DABA. With the identification of the amino acid beta-ODAP as a toxin, it has become possible to select low ODAP genotypes of L. sativus. Such cultivars are now available from Canadian and, to a limited extent, from Indian research programs. Their existence has renewed interest in the further development of this species as a pulse crop. P: Pulse; F: Fodder, forage; O: Ornamental; T: Tubers Lathyrism Genetic resources There are two different types of lathyrism, neurolathyrism and osteolathyrism, affecting the nervous system and bone formation respectively. The term "Human Lathryism syndrome" (HLS or HULAS) was coined to describe the rather diverse clinical and biochemical symptoms (incl. osteolathyrism) caused by L. sativus (Cohn, 1995). The currently ex-situ conserved genetic resources of the genus are mainly restricted to 86 species (FAO, 1993 data). A thorough assessment of the conservation status (ex-situ, in-situ) for the genus and the range of diveristy conserved for individual species is currently being undertaken by IPGRI. In this bibliography particular attention has been given to the inclusion of information about the floras of regions where Lathyrus spp. occur, since floras can contain information about collection sites, habitat, economic botany and the older literature. Although the major utilitarian focus is on Mediterranean Lathyrus species, the genus is no means restricted to this region and others like North-, South-America and Central Asia also offer considerable biodiversity. Cohn and Streifler (1981, 1983) have described, in addition to neurological damage, osteolathyrogenic symptoms in lathyrism patients who had 35 years earlier consumed food prepared from L. sativus seed in a German forced labour camp. This finding suggest that extreme care is needed with the toxicity assessment of low-ODAP strains of L. sativus because other toxins may be present in the seeds under certain conditions e.g. seed immaturity. ii Medical scientists are interested in the causes of neurolathyrism as a model for neurodegenerative diseases striking the more affluent sections of the human population. The scourge of neurolathyrism, known since ancient times, today affects mainly the poorer rural classes especially in India, Bangladesh, Ethiopia, Nepal and Pakistan during drought caused famines. Historic outbreaks of this neurological crippling disease have also been documented for Spain, Algeria, Ukraine, Russia, Germany, Italy, France, Syria and most recently from 1972-74 in China. The cause of neurolathyrism is the continued consumption of L. sativus seed as a staple food. Other Lathyrus species have also occasionally been linked with neurolathyrism e.g. L. cicera, L. ochrus and L. clymenum. These species all contain beta-ODAP in their seeds. on a regular basis. Dr. Kothari in India argues that L. sativus is a safe food and has sparked considerable controversy. Others consider such publicity irresponsible and would rather like to see efforts directed towards better understanding the causes of neurolathyrism. It should not be ignored that malnutrition is an important and constant factor of neurolathyrism epidemiology. A monogastric malnutrition model for neurolathyrism, coupled with detailed research into variation in human ODAP metabolism is likely to provide the best approach to define toxicity levels of this glutamate analogue. In addition, the possibility of other toxins and predisposing factors leading to HLS need to be understood. Without a thorough understanding of the real cause(s) of neurolathyrism and the variability in susceptibility of individuals to it, the bright future of grain crops like L. sativus is handicapped by the stigma of their toxicity. With a thorough knowledge of Lathyrus toxicity and sufficient genetic resources several other species of this diverse genus may well be developed as food crops. Lathyrism and Poverty The occurrence of neurolathyrism is intricately linked to drought caused famine, poverty and malnutrition. The hardy L. sativus may provide most of the food for survival during drought in areas where neurolathyrism is prevalent. Animal fodder Lathyrus fodder crops have been utilised for millennia. It appears that high intake of Lathyrus fodder from BAPN and DABA biosynthesizing species has frequently led to ruminant intoxications. Adaptation by ruminants to such forages has also been reported, whereas horses and pigs can be poisoned by any Lathyrus forage. Lathyrism in horses has been an important military problem in the past. Whether the new low toxin strains will survive under adverse conditons remains to be seen. Detoxification techniques for strains with high beta-ODAP provide some measure to reduce the impact of neurolathyrism, however, as was pointed out by Kearnick and Smartt (1995), the costs in fuel and water may prevent such practise where it is most needed. Recent advances in solar cooking technology through the provision of cheap, easily built solar ovens may reduce the future need for solid and fossil fuels in some regions. Governments need to pay attention to the potential threat of neurolathyrism and provide contingency plans to deal with high risk areas during times of famine. Reduction of poverty and malnutrition in lathyrism prone areas are likely to provide the best solution to this problem. Biomass production by native and introduced legumes has been an incentive for cultivation since the inception of agriculture. Familiarity with naturalised crops and their toxicity has generally resulted in the avoidance of toxic plants and selection of palatable plants for food and feed. Selection amongst existing landraces for adaptation to specific environments and transfer of traditional knowledge about their utilisation provides the basis of new crop options for global agriculture. The development of L. sativus as a safe food crop, not only for Bangladesh, India, Nepal and Ethiopia but also for the peoples of countries with suitable climates to grow this crop (Canada, Chile, Spain, North Africa, Middle East, Afghanistan, Australia etc.) necessitates a good understanding of the conditions and predisposing factors which lead to the onset of neurolathyrism. Models for neurolathyrism and human safety Experimental monogastric animal models for ODAP toxicity are needed. In the absence of suitable protocols to provoke neurolathyrism through oral administration of toxic diets in experimental animals, no safety margin for below-threshold ODAP toxicity can be set. Existing toxicological protocol does not provide for the determination of safe L. sativus consumption levels. Consumption of L. sativus by humans appears to be much safer than previously thought. Large populations in India, Bangladesh and Ethiopia are consume this crop iii improvements can clearly be made in collaboration with others. Collectively, the expertise and knowledge related to the genus Lathyrus could be gathered into a living monograph through a continual reviewing, editing and addition process. Free access to this knowledge depends on whoever controls the copyright. Technical matters Following the work of Davies et al. (1993a) this bibliography was compiled as a database using Procite and Biblio-Link bibliographic software. The aim was to demonstrate the utility of personal bibliographic software (PBS) for the management of subject specific information. This collection provides access to the older literature which cannot be retrieved from current databases. The software facilitates rapid retrieval of information through searches of the database or its indexes. It also allows a range of citation formats to suit even the most discerning editor. Abstracts and references were initially compiled from a wide search for Lathyrus related subject matter using Agricola (1970-1984), CAB (1984- June1994), BIOSIS (1991- June 1993) and Medline (1966-1993) CD-ROM data spanning the period 1970-1993. Search strategy: Lathyrus or grasspea or (grass and pea) or chickling or lathyrism. These records were imported into Procite 2.02 with the help of the bibliolink 1.1 software for silverplatter. Extensive editing was required to achieve a homogeneous format. This computerised search was complemented by a thorough check of Herbage Abstracts (HA) for Lathyrus back to the year 1931. Additional citations were obtained from the Lathyrus literature and for taxonomic references from the USDA pcGRIN database for Lathyrus [Lathyrus.exe]. ICARDA kindly provided additional entries from CODIS (1975-1995). Some references were obtained from Dissertation abstracts and the most recent citations originate from Current contents. The more recent version of BiblioLink® II was used to import these records with ease and flexibility thus necessitating only minor editing. Spell checking of the database was achieved by exporting the records to a text file and then importing this file in MSWord 6.0. This bibliography provides a thorough coverage of the agricultural, botanical, chemical, biochemical and medicinal literature related to the genus Lathyrus and neurolathyrism. Every attempt has been made to make this collection as complete as possible. It is, however, just the start for future additions. It is hoped that others will join in the effort to make future editions more comprehensive. In order to assist researchers and other interested individuals to utilise the collected information, CLIMA and ICARDA have agreed to publish this bibliography in both electronic format (CD-ROM) and as a hard copy. With such a comprehensive bibliography the foundations are laid for a truly encyclopedic database for the genus Lathyrus. It is perceivable that in the near future, books are to be published primarily in electronic format. Monographs may take on the shape and form of encyclopedic multimedia databases, encompassing all available knowledge on a given topic. Revisions, discussions and updates of particular areas can be part of an ongoing process. A book needs to be reprinted for every update but a database monograph can be continually updated thus providing a comprehensive, up-to-date and unified body of knowledge on the subject. Provision of collection points for individual contributions in the form of bibliographies such as this one, for information which currently is not freely accessible by computer, facilitates the creation of ever more complete knowledge databases. It is hoped that the present work provides a stimulus for similar activity with other crops and research interests. A bibliographic database like the present one on the topic Lathyrus can be crosslinked to data and full text publications, archives, genetic resource inventories, evaluation data, maps and models, spatially referenced data etc. With publications in CD-ROM format and the vast possibilities of the internet, a great deal of storage space is available for other information to be added in future editions. Subject indexes An attempt has been made to index each record with a minimum of 3 key words to provide several independent access routes to it. In many records the format “species”, “species topic” and “topic species” has been used to provide three separate index entries, so that records do not get lost in the database. Species occurrence is noted, especially for floras, and sometimes includes synonyms. All Lathyrus species in the index were searched for individually in the database. Taxonomic indexing is useful for the retrieval of information on the rarer species, it also provides an index of the lack of knowledge about these lesser known species of the genus. Some references (mainly those extracted from pcGRIN) appear in abbreviated notation which should be familiar to taxonomic botanist. Species names indexed with their authority name refer to publications where they were authoritatively described. Further details about the taxonomic literature of the genus Lathyrus can be obtained from ILDIS. Bibliographic databases as the nucleus for assembling the body of knowledge on a given topic Publication of a printout from the database necessitated the creation of a subject index in order to facilitate manual location of appropriate records. For the subject A PC database format facilitates a mini expert system. Such a system is only as good as its programmer, so iv Libraries index, full text searches of the bibliographic database using major topics and each legume or crop species were carried out to facilitate manual retrieval of records from the hard copy. The index terms are of heterogeneous origin, since a fair portion of the records in this database originated from other databases e.g Agricola, CAB, BIOSIS., AGRIS and Medline. Additional index terms were created during the manual addition of records from Herbage Abstracts and literature citations. Some overlap occurred as a consequence. Plant species are indexed by their latin binomials. Legume species were included in the index when mentioned in the abstract, in order to provide linking terms with any future bibliographies. The following libraries with extensive collections were visited or consulted for this bibliography: Agricultural library Stuttgart, Hohenheim Agricultural library, Berlin Barr Smith Library, University of Adelaide Bibliothek der Tierärztlichen Hochschule, Hannover, Germany Biological Sciences Library, University of Western Australia, Perth, Centro de Ciencias medio-Ambientales, Serrano, 115 Bis., Madrid, Spain David Lubin Memorial Library, FAO, Rome ENSA pasture library, Montpellier Harold Woolhouse library, Waite Agricultural Research Institute, University of Adelaide, South Australia; ICARDA, Aleppo, Syria Instituto Nacional de Investigaciones Agrarias, Ctra. De la Coruna, 7, Madrid, Spain ISIS http://www.nalusda.gov/isis/ Library of the Adelaide Botanical Gardens (Herbals, Floras) Library of the Ibero-American Institute, Potsdamer Platz, Berlin, Germany (Spanish literature) Library of the IPK, Gatersleben Library of the Royal Botanic gardens, Kew, Great Britain Rothamstedt library, Harpenden, Great Britain Staatsbibliothek Zu Berlin -Preussischer Kulturbesitz, Potsdamer Platz, Berlin, Germany USDA National Agricultural library (NAL) Zentralbibliothek der Landbauwissenschaft und Abteilungsbibliothek für Naturwissenschaft und vorklinische Medizin, Bonn, Germany The staff of these institutions are gratefully acknowledged for their help with the obtention of materials for the reprint collection. It is possible that some records may have been indexed erroneously. Several topics have been indexed using a combination of index terms thus increasing the possibility of mistakes. Seemingly unrelated subject links were included where it felt appropriate to draw interested readers to a particular subject e.g. poverty, socioeconomics and malnutrition. Reprint collections Records for which reprints have been obtained are labelled as such (Reprint= CLIMA collection, reprintDE= Dirk Enneking collection). The location and assembly of a reprint collection for the remainder of the literature is a formidable task and may take decades to accomplish. A safe depository or archive for this collection also needs to be found to guarantee access for future generations. Receival of reprints and photocopies, especially copies of articles from the older literature and the rarer journals would be very welcome. Other reprint collections should be documented and noted in updates of the database to facilitate access to literature. Acknowledgments The following people helped in one way or another and their contribution is gratefully acknowledged: Christine Davies, Dr. Ann Butler, Dr. Ann-Marie Arentoft, Dr. K. H. M. Siddique, Dr. Ashutosh Sarker, Dr. Clayton Campbell, Dr. D. N. Roy, Dr. John Hamblin, Ian Maling, Dr. M. C. Saxena, Dr. L. Robertson, Dr. S. Varma, Nihad Maliha, Dr. Max E. Tate, Ken Street, Dr. Y.-H. Kuo, Dr. N. K.. Narayan, Prof. Peter Spencer, Dr. C. Assmussen, Dr. H. K. M. Yusuf, Prof. S. L. Mehta, Tatiana Alexandrova and IPGRI. I would like to thank Dr. Clive M. Francis, Prof. Lambein and Ken Street for their critical comments on the introduction, and especially Mike W. Perry and Prof. Fernand Lambein for their encouragement to persevere with this task. Copyright Considerations Reprints are noted and ref. sources acknowledged, either by the record numbers of the major databases or by author name and year. Obtained permissions to print copyright material is acknowledged at the bottom of the abstract, prior to the index terms. Permission to publish abstracts was sought from publishers of the respective journals. The response to these requests was very positive and is gratefully acknowledged (see section “Copyright permissions” below) Abstracts obtained from CAB and BIOSIS (noted in the code number field) are licensed on the basis or the number of abstracts/copy of the bibliography. The costs incurred are included in the purchase price of this bibliography v Copyright permissions The following publishers have kindly provided permission to publish abstracts from the journals listed Publishers Academic Press: Analytical Biochemistry, Arch. Biochem. Biophys, Environmental Research, Fundamental and Applied Toxicology, Toxicology and Applied Pharmacology Munksgaard Publishers, Copenhagen: Acta Neurologica Scandinavica American Chemical Society: Biochemistry American College of Veterinary Toxicologists: Veterinary and Human Toxicology American Journal of Tropical Medicine and Hygiene: American Journal of Tropical Medicine and Hygiene American Society of Biological Chemists Inc., USA: Journal of Biological Chemistry Blackwell Science Ltd.: Heredity, Journal of Ecology Carfax Publishing Company, UK: British Poultry Science Cytologia, Tokyo: Cytologia, Tokyo Dr. W. Junk Publishers, Netherlands: Genetica, Qualitas Plantarum/Plant Foods for Human Nutrition Elsevier Science B.V., Amsterdam, Netherlands: Biochimica et Biophysica Acta, Brain Research, FEBS Letters, Immunology Letters Elsevier Science Ireland Ltd.: Neuroscience Letters, Toxicology Letters (Amsterdam) Elsevier Science Ltd., The Boulevard, Langford Lane, Kidlington 0X5 1GB, UK: Phytochemistry, Soil Biology and Biochemistry Ethiopian Medical Journal: Ethiopian Medical Journal Folia Histochem Cytochem Krakow: Folia Histochemica et Cytobiologica Gustav Fischer Verlag, Stuttgart: Zentralbl Bakteriol Naturwiss International Society for Horticultural Science: Acta Horticulturae Kluwer Academic Publishers, Netherlands: Euphytica Macmillian Journals Ltd, UK: Brain Marcel Dekker Inc.: Communications in Soil Science and Plant Analysis, Preparative Biochemistry Masson S.A. Paris, France: Bull. Soc Pathol Exot Filiales Raven Press: Journal of Neurochemistry S Karger AG, Basel, Switzerland: European Neurology Springer Verlag: Acta Neuropathol Berl, Arch Toxicol Supplement, European Journal of Biochemistry, Experimental Brain Research, Theoretical and Applied Genetics Stockton Press: Paraplegia The Biochemical Society: Biochemical Journal Walter de Gruyter & Co., Berlin: Biol Chem Hoppe Seyler Journals Journal Acta Horticulturae Acta Neurologica Scandinavica Acta Neuropathol Berl American Journal of Tropical Medicine and Hygiene Analytical Biochemistry Arch Toxicol Supplement Arch. Biochem. Biophys, Biochemical Journal Biochemistry Biochimica et Biophysica Acta Biol Chem Hoppe Seyler Brain Brain Research British Poultry Science Bull. Soc Pathol Exot Filiales Communications in Soil Science and Plant Analysis Cytologia, Tokyo Environmental Research Ethiopian Medical Journal Euphytica European Journal of Biochemistry European Neurology Experimental Brain Research FEBS Letters Folia Histochemica et Cytobiologica Fundamental and Applied Toxicology Genetica Heredity Immunology Letters Journal of Biological Chemistry Journal of Ecology Journal of Neurochemistry Neuroscience Letters Paraplegia Phytochemistry Preparative Biochemistry Qualitas Plantarum/Plant Foods for Human Nutrition Soil Biology and Biochemistry Theoretical and Applied Genetics Toxicology and Applied Pharmacology Toxicology Letters (Amsterdam) Veterinary and Human Toxicology Zentralbl Bakteriol Naturwiss Publisher International Society for Horticultural Science Munksgaard Publishers Springer Verlag American Journal of Tropical Medicine and Hygiene Academic Press Springer Verlag Academic Press The Biochemical Society American Chemical Society Elsevier Science B.V., Amsterdam, Netherlands Walter de Gruyter & Co., Berlin Macmillian Journals Ltd, UK Elsevier Science B.V., Amsterdam, Netherlands Carfax Publishing Company, UK Masson S.A. Paris, France Marcel Dekker Inc Cytologia, Tokyo Academic Press Ethiopian Medical Journal Kluwer Academic Publishers, Netherlands Springer Verlag S Karger AG, Basel, Switzerland Springer Verlag Elsevier Science B.V., Amsterdam, Netherlands Folia Histochem Cytochem Krakow Academic Press Dr. W. Junk Publishers, Netherlands Blackwell Science Ltd. Elsevier Science B.V., Amsterdam, Netherlands American Society of Biological Chemists Inc., USA Blackwell Science Ltd. Raven Press Elsevier Science Ireland Ltd. Stockton Press Elsevier Science Ltd., The Boulevard, Langford Lane Marcel Dekker Inc Dr. W. Junk Publishers, Netherlands Elsevier Science Ltd., The Boulevard, Langford Lane Springer Verlag Academic Press Elsevier Science Ireland Ltd. American College of Veterinary Toxicologists Gustav Fischer Verlag, Stuttgart 6 Abbreviations Annotations are initialled by author. Agricola AGRIS BB BIOSIS BOAA CA CAB CD-ROM CGC_95_2 CLIMA CP Cv D.E. DABA DM ENSA FAO FL HA i.v. ICARDA IPGRI ISIS LWB M.E.T. Medline NAL NPAA UWA ODAP PBS PC pcGRIN TRANSL UB USDA USDA agricultural literature database FAO agricultural literature database BeanBag BiosSiences Information Services (Biological abstracts) beta-oxalyl-amino-alanine (=ODAP) Chemical Abstracts Commonwealth Agricultural Bureau Computer Disc-Read Only Memory references added by Clayton G. Campbell February 1995 Centre for Legumes in Mediterranean Agriculture Crude protein Cultivar Dirk Enneking comments Diamino butyric acid Dry matter Ecole Nationale Superieur d'Agricole Food and Agriculture Organization of the United Nations Fernleihe=Interlibrary loan Herbage Abstracts intravenous injection International Centre for Agricultural Research in Dry Areas International Plant Genetic Resources Institute Integrated system for information services Landwirtschaftliche Bibliothek, Bonn Max E. Tate comments Medical abstract information service National Agricultural library, USDA Non protein amino acid University of Western Australia Oxalyl-diamino-propionic acid Personal bibliographic software Personal computer Personal Computer Genetic Resource Information System Translation Unibibliothek-Adenauer Allee, Bonn United States Department of Agriculture 7 Table 2. Gen L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L Species alpestris amphicarpos angulatus annuus aphaca arizonicus armenus atropatanus aureus basalticus bauhinii bertianus bijugatus binatus blepharicarpus boissieri brachycalyx brachypterus cabrerianus campestris cassius chloranthus chrysanthus cicera cilicicus ciliolatus cirrhosus clymenum cyaneus czeczottianus davidii delnorticus dielstanus digitatus dominianus elongatus emodi eucosmus filiformis frolovii gloeospermus gmelinii gorgoni graminifolius grandiflorus hallersteinii hasslerianus heterophyllus hierosolymitanus hirsutus hirticarpus hitchcockianus holochlorus hookeri humilis hygrophilus inconspicuus incurvus japonicus jepsonii karsianus komarovii krylovii laetiflorus laevigatus lanszwertii latifolius laxiflorus layardii ledebouri lentiformis leucanthus libani linearifolius linifolius littoralis Author (Waldst. & Kit.) Kit. L. L. L. L. Britton (Boiss. & Huet) Celak (Grossh.) Sirj. (Steven) Brandza Rech. F. Genty Colla T. White Pancic Boiss. Sirj. Rydb. Celak. Burkart Philippi Boiss. Boiss. Boiss. L. Hayek & Siehe Rech. F. Ser. L. (Steven) K. Koch Baessler Hance C. Hitchc. Harms (M. Bieb.) Fiori Litv. (Bornm.) Sirj. Fritsch Butters & St. John (Lam.) Gay Rupr. Warb. & Eig. Fritsch Parl. (S. Watson) T. White Sibth. & Smith Baumg. Burkart L. Boiss. L. Mattatia & Heyn Barneby & Reveal (Piper) C. Hitchc. G. Don (Ser.) Sprengel Taubert L. (Roth) Willd. Willd. E. Greene P. H. Davis Ohwi C. Serg. E. Greene (Waldst. & Kit.) Gren. Kellogg L. (Desf.) Kuntze Ball ex Boiss. Trautv. Plitm. Rydb. Fritsch Vogel (Reichard) Baessler (Nutt.) Endl. The genus Lathyrus (Kupicha, 1976) Section Orobus Lathyrus Linearicarpus Lathyrus Aphaca Orobus Lathyrostylis Lathyrostylis Orobus Lathyrus Lathyrostylis NotoLathyrus Orobus Pratensis Lathyrus Lathyrostylis Orobus Lathyrostylis NotoLathyrus NotoLathyrus Lathyrus Lathyrus Lathyrus Lathyrus Lathyrostylis Lathyrus Lathyrus Clymenum Lathyrostylis Pratensis Orobus Orobus Orobus Lathyrostylis Orobus Lathyrostylis Orobus Orobus Lathyrostylis Orobus Clymenum Orobus Lathyrus Orobus Lathyrus Pratensis NotoLathyrus Lathyrus Lathyrus Lathyrus Lathyrus Orobus Orobus NotoLathyrus Orobus Linearicarpus Linearicarpus Orobus Orobus Orobus Lathyrostylis Orobus Orobus Orobus Orobus Orobus Lathyrus Pratensis Pratensis Lathyrostylis Lathyrus Orobus Orobus NotoLathyrus Orobus Orobus Gen L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L L 8 Species lomanus longipes lycicus macropus macrostachys marmoratus mulkak multiceps nervosus neurolobus nevadensis niger nigrivalvis nissolia nivalis occidentalis ochroleucus ochrus odoratus pallescens palustris pancicii pannonicus paraguayensis paranensis parodii parvifolius pauciflorus phaselitanus pisiformis polymorphus polyphyllus pratensis pseudo-cicera pubescens pusillus quinquenervus rigidus roseus rotundifolius satdaghensis sativus saxatilis setifolius spathulatus sphaericus splendens stenolobus stenophyllus subandinus subulatus sulphureus sylvestris tauricola tingitanus tomentosus torreyi trachycarpus tracyi transsilvanicus tremolsianus tropicalandinus tuberosus tukhtensis undulatus vaniottii variabilis venetus venosus vernus vestitus vinealis whitei wilsonii woronowii zionis Author I. M. Johnston Philippi Boiss. Gillies Vogel Boiss. & Blanche Lipsky D. Clos Lam. Boiss. & Heldr. S. Watson (L.) Bernh. Burkart L. Hand.-Mazz. (Fischer & Meyer) Fritsch Hook. f. (L.) DC L. (M. Bieb.) K. Koch L. (Jurisic) Adamovic (Jacq.) Garcke Hassler Burkart Burkart S. Watson Fern. Huber-Mor. & P. H. Davis L. Nutt. Nutt. L. Pampan. Hook. & Arn. Elliott (Miq.) Litv. T. White Steven Willd. P. H. Davis L. (Vent.) Vis. L. Celak. Retz. Kellogg Boiss. Boiss. & Heldr. Philippi Lam. Brewer L. P. H. Davis L. Lam. A. Gray (Boiss.) Boiss. Bradshaw (Sprengel) Reichb. f. Pau Burkart L. Czeczott Boiss. Leveille (Boiss. & Kotschy) Celak. (Miller) Wohlf. Muehlenb. (L.) Bernh. Nutt. Boiss. & Noe Kupicha Craib Bornm. C. Hitchc. Section NotoLathyrus NotoLathyrus Lathyrus NotoLathyrus NotoLathyrus Lathyrus Lathyrus NotoLathyrus NotoLathyrus Neurolobus Orobus Orobus NotoLathyrus Nissolia Lathyrostylis Orobus Orobus Clymenum Lathyrus Lathyrostylis Orobus Lathyrostylis Lathyrostylis NotoLathyrus NotoLathyrus NotoLathyrus Orobus Orobus Lathyrus Orobus Orobus Orobus Pratensis Lathyrus NotoLathyrus NotoLathyrus Orobus Orobus Orobon Lathyrus Lathyrostylis Lathyrus Viciopsis Orobastrum Lathyrostylis Linearicarpus Orobus Aphaca Lathyrus NotoLathyrus NotoLathyrus Orobus Lathyrus Linearicarpus Lathyrus NotoLathyrus Orobus Lathyrus Orobus Orobus Lathyrus NotoLathyrus Lathyrus Lathyrostylis Lathyrus Orobus Lathyrostylis Orobus Orobus Orobus Orobus Linearicarpus Orobus Orobus Linearicarpus Orobus