THE LICHEN GENUS NEPHROMA IN. NORTf{ AND l"'J:DDl,E AMER!GA by Ollft"ord Major Wetm$·re AN ABSTRAOT Submitted to the Gollege of So1ence and Arts ot Michigan State Un1vera1tr ot ~1oalture and APplled Science 1n partial t'Ulf1llment c;;t the requlrfments ro:r the degree ot MAS'BB OF SOIENCE Oepartment of Botany and Plant' Pathology 19.59 a- ~ · ~+ The morpholog1CJti:1 and anatomleal teat.u•es are tl.lso~tHaed !n detail and twenty-nine original figures re 1n¢1 Q.dttd. The protuberance& on the lower su.rf·ace ot !• r:ts~p1.na1\um {L.) Ach. are eortioate and, thet-erore, not tx-ue peeudo-eyphellae as often recorded 1n the literatul"e• Eight 11ohert subs~anoes have been fcrun4 1n the genua: ~ante ae1d; nephrom1n• ze.or1n; nephl'in and four· un1dentif'1ed Jteutral au't>stances. i.t'hfHie 1 have taxonom.!.c although other wo:-ken tubs~noes S.mpo~tanee, 1n r-ela tea ge.ne~ have ignored them. ~e genus 1s Ple.oed in the Nephromae·e$e instead of the lih•l'e uauaJ..ly ot the ~;~~·.,.,,,..,.._ ~las·e1tled; b-~au.tu~ i•_(; ~~~i.!rfl.J.l.i)-tt8 1n a.~u4 etruotuJte and de:'rele>-pment; conidial p~ 4u~,~'t.G,h .ud the pJ-e.setloe of a lowe~ c.ol'tex 1n N~Rhtsm.• ~ Seven epeo1es and two eubspeeies are t-eoogf.\1zed ·and a ~;e~··:to~ thetr 1denttt1cat1.on 1s 1neluded along with a . .!·.· ": cU.agitos1.• • n~merun.e.tttraJ. notes, tlnd a cl1atrlbutton map tor species. ea~h Gyeln1lt•s rour new speo1ee and two new 1nfraapeo1flc taxa t'irom Nor·th Atnerioa are reduoed to syn:onymy. N;etph£t.!Pa trop1own (MiiJ.l Arg.) Zahlb. i s reduced to a subspecies of ' !!· hel vet1cum Ach. Nephroma bellum (Spren.g.) ',tuck. is recognized as the correct name for !!· l$.ev1gatum s uet. non Ao'h. and 11· laev1g&tum Aeh. non auct. 1s recognized as the correct name for what previously has be.en called ! · 11.1&1 tan Scbae:r. i.e_~ The need for tutu.re work 1n the ohemletry, algal and oul. tq:re, and phylegeny 1s reoogntted together with the spor~ need tor study or the remaining speo1es of Nephroma in the world. THE LIOJIEN GENUS NEl?fiROM,A. IN NORTH A!Ul I<· IDDLlt AMERICA by Major Wetmore Gl1ffo~ A THESIS or or to the Gollege Sci-ence an4 Arts Su,bm1t·t~d MiolU.gan State t1n1vet-s1 ty or AgrtoUl tHa.re and Appll.ed Se1eaoe in partial rtafillment or tbe requtrements tor the ot deg~ee MASfER OF SCIENCE »e.partlllent or Botany and Plant .PatbolOSJ' 1959 P ge LlSTING f>Jt PLATES • • 1 I . INTRODUCTION • • • • • • • • • • • • 1 II. • • • • • • • • • • • • • • • • • • 3 III. MORPHOLOGY • • • • • • • • 6 A. Methoda • • • • • • • • • • • • • • • • • 6 D. fhe.llus • • • • • • • • • • • • • • 8 o. Apothee1 • • • • • • • • • • • 12 • e • • D. Pyen1d1a. • • • • • • • • • • • • • 14 IV. CHEMISTRY • • • • • • • • • • • • • • • • 16 A. Methode • • • • • • • • • • • • • • • • 16 B. Sub tanoes • • • • • • • • • • • • • • • 18 v. ECOLOGY • • • • • • • • • • • • • • • • • • • 24 VI. DISTlUBt1l'!ON • • • • • • • • • • • • 26 VII. TAX-oNOMY • • • • • • • • • • • • 27 A. Family • • • • • • • • 27 8. Genus • • • • • • • • • • • • • • • 28 c. Key to species • • • • • • • • • • • • 29 l. tu;gt1cum • • • • • 30 2. expalll<1 \Ill • • • • • • • • • • • 33 ) . :re~_HRtAAtum • • • • • • • • • • • J4 4. !!• beJ,lum • • • • • • • • • • • • 37 '· !i• pelyet1oum • • • • • • • • • • • ~ LISTING OF PLATES Plate I. Detailed drawings of oephalod1a of ~· atcti~tU! and !· ex;pa.llidum. Plate II. DetaUed drawings of cortex, algal layer Md med uJ.la o t Nephroma. Plate III. Detailed cross eeotional draw1nge of sored1a and margin of an apotheouem. ~~e Plate IV. Draw~ngs of oross section of a papilla. of !i• or resHU!.nat·Ull'l and a ps- udocyphella Pteudootph fl..le.r,la. v. Plate Dntwinge of tips of aeo1 from Pe.Jt~gem, SOl.ottina, ·and N Rh.roma. After Gal1nou 195,. Plate VI. Drawings of nol"'Illll spores of each species of Nephroma and ot abnormal and germ1na t!ng spor.es. Plate VII. Detailed drawings of conidlophores of 1:!· he}. v~t1oum. • Pl'ate VI!I. Photographs of oryetale of neutral substances recrystallized 1n G.A.An. Plate IX. Photographs of crystals ot teor1n, nephr1n and carotenoid recrystallized in G.A.An. or Plate X. Detailed drawings tomentum on lower eurfaee ot thallus a.nd of medUllary hyphae. !!.· Plate XI. Dietr1but1on map of a~-ot1.0YJ!• l te Xll. D1str1bution map of N. eJtP!llldum. 1 te XIII. Distribution map of H· tei!e1n~tum. te XIV. Distribution map ot li· beJ.J.wn. . I xv. te D1str1but1on map of N. h•lvetioum. ...-,. i -·;, t t te XVI. Distributton map of !!· 1 ev\satum, . ~ . I te XVII. Distribution map of !!,~ Mr~J.,e. 1 'I' I Dl'PB.ODUO TION This. 1 a revisionary tr~atment of the genua N_epht'ol'l\a, a g nu of 11ohen1z :'.. A eomyc te of the D1soomy-cete group, 1n the ~lorthern Hemisphere of the t ett World, including Gr e - land on the e t , .Alaek on the west, and Pan~UDA on t..l-ie south. In thi study I h ve exam1n 1 ,918 pee1men from 16 1n t1tu tional herb ri in th Un 1 t d Ste. tea and Can a a a d .5 pr1- vate he~bar1a~l The herbarium mater1 wa upplemented with fisld studies and collections in !sl Royale National Park (Michigan) , and in Haiti an Dominican Republic 1n th Caribbean. Fro-a this field study, l was able to observe the n tural \l'ari t1on, habitat, and o ourrence, as well as make 1 rge collections of material for later study. J -L1 t ot herbar1$. Abbr e\l'ia t1on fo·ll<DW the recom- ndat1QnS of Lanjow and St fleu (1959). t.1ollen Herb., Am. ol. So~. (ABSL}; Nat. F~gu.s Colla., Hort. Crops Reaearoh nob ( BPI ) ; Na. t . Mus. Canada ( c:AN ) ; Un1 v. of Cpl. ( COLO ) ; 1o go Mat. JU•t. Mus.. (F); farlow Herbarium, He.rvara Unlv. H); Uh1 v. ot 1ch. (MIOM); M1ssou.r1 Bot. Gard.. ( MO ); Mich. t Univ. (MSO); l4ew York Bot. Ga.rd. (NY); Phil d·elph1 4. ot Nat. So1. (#>H); t.Tn1v. of Toronto (TRTO); un1v. ot lit. (UO)j Smithsonian Inst. (US); \ielleeley Coll. (WELC); ..-. of 9'1Jo. ( IS). I am gratefUl to the curators ot a he:tba:r1a tor the loan of this material. \' (:IN' t ' • ~~ Tne 1t.ollow1ng personal herbaria were kindly lo ned and o. 1thout abbreViation • ttl. L. CUlberson (CUlberson), W. w. (Dodge), • E:. Hale {Hale), F. P. S1pe (S1pe) and J . on ( Thomeon). 1 2 This is not monograph beeause of 11m1 tat ions in scope &. necessitated by t1me and faoU1t1es. However, I have a.t tempte.d. to examine e.nd r-e-evaluate as many ot the characters. that he.ve been J>eported in the. literature a.a possible, to A reaffirm thelr presence and to eva~ t.Utte the1r toxonom1e II HISTORY t ,I-" (. . ~.r' r'lt: L1nnaeus (1?$3) plaoed the two species 9f Nep9£2ma -- - I kn.e>wn at his t1me..... _)1n his genus L1ohen. which included all speoies wh1oh he ooneidered lichens. He reQogn1zed J...• arct1cus and t:.• 4es,.u'iina.tus. - . ( . $ • ' In 1794 iehrader included. reaup1nata in the genua ~!31 tlgem tor the fit"st time. He only ~eported t• re.sup1na.ta and f· !:[lvatic.a ·in. this genus 1n thi~ paper• ...! ./ ·-t- I '\ / I In 1803 Aehariu.s placed. the :reoogn1zed • speq1e&~:t' 1 ** Nep~roma) into e. separate seet1on of Pel.t1sera (Pel tis}~ In 1610 he recognized this eeetion as a separate Op1ste~1a). genus and changed the name to Nephto!p.a. Aeharius reeogn1zed • poler1p, H_. ~esq;e-ina..:t:f, !· I?a~111s., and li· ~:t,vetlea.. i'llt-:4!' lit ,Aohar1us used the natlle in the feminine ne in N~ph;:oma error, for he states that the tef'm came from the Greek word r kidney ( 4.n reference to the apotneo1sl shape), hloh wa_!) 7 vl· ,·• ~ r/< ~.,.a_ ~~,_,..o .. _. ) - --- otually neuter. This error of gender was corrected in 1812 11 5. F. Gray to the neuter gender. ln 1860 Nylander deser1bed the genus Neph;r:oma.um for apeo1es ot flephrqm~. with gon1m1a (blue green algae) and Nephroma for those species w1 th gon1d1a (green HoweveT, he apparently considered the blue green 1n the eephalod1a of ! · ext:eJ-l14wn to be ot sufficient . I 3 4 //....-.... importance to plaoe \!-~1;~ 1n h1a genus N~phromium. In 1909 Wainio raised Aohar1ue' sect1on _Op1ster1a to generic rank oh the 1den thAt the oldest ns.me_ regardless of rank, had PJ>1Cr1ty, 1e oontre.ry to the present rules ot w~ioh nomenclature (Lanjouw, 1956). Since Nephroma was the first name used for these spe0ies in the rank of genus, it has priority over Qpis.ter.ta. • The next work et importance 1n the genus was dorie by Moreau (1927) when he p~oposed the family Nephrom1aoees for the genua His for the sep ration from H!phro~A~.. arg~ent the Peltlgeraceae was based on the faet that has a Nephr<?lll!~_ lower cortex and does not. This separation was ~eltige~a later suPPorted by cytalog1ca.l differences in aeoogon1e.l development (Moreau, 1928) and difference a 1n the apex of the scus (Ge.J.inou, 1955). 'l'he only monogt-a:ph1c attempt of fie.ohroms. w: e done in I the 1930's by Gyeln!k (1930, 1931, 19J2a, 19J2b, 1932c, 1934, 935, 1936, s.nd 1940) in Budapest. His ms1n species raotere l<rere the degree of tomentum, KOH reaot1ons, lus color, eoredia, and tubercles. By using every small t1on and combination of these, and a f ew other characters, oribed 28 new species for the Northern Hemisphere, 4 of ere from North. America, and 45 infraepeo1fic names, trom North America.. Most of these taxa have not been 7 . l;r-<-:t~ ..., ,tl!li',1zed by other 11chenolog1sts. Gyel nik also worked 1n . . ~- oth r groups, but mostly in the Peltigeraceae. {1954) state& that of the 1,307 new names described by , only a few are accepted today. However, almost all