THE RECENT CRINOIDS OF THE COASTS OF AFRICA.
By Austin Hobart Clark,
AssistantCurator,Division ofMarineInvertebrates, U.S. NationalMuseum.
HISTORY.
The
history of the study of the recent crinoids inhabiting the coasts of Africa can be stated in a few words.With
but three exceptions all the references to African crinoids are only incidental incorporatedinworks sometimes
restricted to the echinodermsalone, butmore commonly
very general in scope.Thus
while the titles of booksand
papersnumber
aboutseventy,from
each oneonly a smallamount
of information is gained,and
an adequate conception of African crinoids as a whole can be acquired onlyby
anamount
of labor totallyincommensurate
with thesum
of theknowledge
gained.The
first crinoidknown from
African waterswas
describedfrom
Mauritius in 1816by Lamarck under
thename
of Comatula carinata.Lamarck
adopted thename
carinatafrom
Leach,who
inthe previous yearhad
diagnosed, inaveryinsufficientmanner,
hisAlectocarinata,which Lamarck
thoughtmight
turn out tobe his species.In 1817 the portion of Savigny's Description de l'Egypte dealing with the echinoderms
was
published,and
in it were figuredtwo
comatulidsfrom
theRed
Sea, one ofwhich was
designatedby Audouin
as"
Comatula sp.," the other as"
Comatula rnultiradiata."There is
no
further reference to the first of these figures,which
rep- resents Tropiometra encrinus; but in 1836 de Blainville copied the second in the atlas to hisManuel
d'Actinologie. In doing this hemade
a curious mistake, for the plate islettered"Comatula
adeonse,"though in the text the description of Comatula adeonxis taken
from Lamarck, and
thespecies is correctly said tohave
ten arms. In the followingyear thePenny
EncyclopediacopieddeBlainville'saccount of Comatula adeonx, multiradiate figureand
all,and
thesame
slipwas made by Knight
in his NaturalHistory, publishedin 1867.Riippel,inthecourseof his travels,found in the
Red
Sea an inter- esting multiradiate comatulidupon which
he bestowed themanu-
Proceedinqs U.S. National Museum,Vol.40—No. 1808.
80796°—Proc.N.M.vol.40—
11—
1
1
PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.script
name
of Comatula leucomelas, but Ihave
notbeen able to find that hementioned
itanywhere
in his works. In 1833 Leuckartcame
across his specimens in theSenckenberg Museum
at Frankforton Main and
published thename,
together with the locality,though
withoutany
diagnosis.Leuckart
was
the first to describe the curious parasiticworms
belonging to the genusMyzostoma
withwhich
crinoids are usually infested, his attention having beenfirst called tothem by
mistaking one for a madreporic plate. In discussing the genusMyzostoma
he mentions a multiradiate comatulidfrom
theRed
Sea, which, follow- ing Audouin, he identifies as Comatula multiradiata,but which von
Graff, acting
on
the advice of P.H.
Carpenter, has suggestedwas
probablyan example
of Heterometra savignii, the species towhich
Audouin's Comatula multiradiatahas alwaysbeen
referred.In theIconographie
du Regne
Animal, publishedby
Guerin-Mene-ville
from
1828-1837, there aretwo
figures1 supposed to represent the species described as Comatula carinatafrom
Mauritius. Possibly the first does represent this species,though
it looksmore
likesome
species of Antedon;
but
the second (2a) appears to be a species of AmpTiimetra,and
agrees fairly well with A. discoideafrom
northern Australiaand
theEast
Indies.There
is a specimen of Ampliimetra discoidea (labeledby
P.H.
CarpenterAntedon
milberti, var. dibra- chiata) in the ParisMuseum from which
I suspect the figurewas
drawn.In 1841
Johannes
Muller described his Alecto savignii,which was
basedupon
specimenswhich had been
broughtfrom
theRed
Seaby Hempricht and
Ehrenberg,now
in the BerlinMuseum, and
he also identified Savigny's figure,which had
been called Comatula multi- radiataby
Audouin, as this species.Two
years later he described Alecto wahlbergiifrom
specimens broughtby Wahlberg from
Port Natal,which
heexamined
at theStockholm Museum.
Michelin in 1845 noted the occurrence of Comatula carinata at Mauritius.
His
specimens arenow
in the ParisMuseum.
In 1849 the comprehensive
monograph
of Muller, completing his studieson
the comatulids, gave asummary
oftheknowledge
inregard to African species at that date. Practically thesame
accountwas
given in 1862by
Dujardinand Hupe
in theirmonograph on
the so-calledZoophytes.Bohlsche in 1866 described as
new
a curious little comatulidfrom
the coast of Brazilwhich
hehad been
unable to identify withany
previouslyknown
species.He
called it, incompliment
to thejustly
famous Norwegian
naturalist of thatname, Antedon
dubenii.This species has
been
the cause of considerable confusion. P.H.
iZoophytes,pi.1, figs. 2,2a.
no.1S08.
RECENT AFRICAN CRINOIDS— CLARK.
3 Carpenteridentified withit a specimenwhich
the Challenger dredged atBahia,and
figuredboththisspecimenand
thetypeinthe Cliallenger reporton
the Comatulse.The
Cliallengerspecimenis ayoung exam-
ple of Tropiometra picta, butthe type-specimen obviously belongs to the Antedonidse
and
to the genus Antedon.Nothing
like it has since beenfoundon
theAmerican
side ofthe Atlantic.E.
Von
Martens in 1869 recordedfrom
theRed
Sea the Alecto palmata of Miiller,which had
originally been describedfrom
India,and
at thesame
time recorded ComatulaSolarisfrom
Zanzibar.The
determination of the formerwas
correct, but the latter appears tohave
beenin reality Tropiometracarinata.Von
Martens didnot dis- cover that the species recordedby
Leuckart as Comatula leucomelasis the
same
as the one givenby
himself as Comatula palmata.Sir C.Wyville
Thomson,
in hispreliminary reportupon
thecrinoids of thePorcupine expeditionand
in his semipopularwork The Depths
of the Sea, publishedin 1873, as wellas in
The
Atlantic, publishedin 1877, touchesupon
the fauna of the Mediterranean, but the only crinoid he mentionsfrom
the vicinity of Africa is " Rhizocrinus lojfotensis,"which was
dredgedby
the Swedish frigate Josephineon
theJosephineBank.
In 1878 Pourtales
and
in 1879Rathbun
discussed at considerable length specimens of Tropiometra carinatafrom
Zanzibar,comparing them
with specimensof T. pictafrom
the coast ofBrazil.In 1879 also
Edgar
A.Smith
described in detail anew
comatulid, Comatulaindica,from
theisland of Rodriguez,which
remains to-day theonly crinoidknown from
that locality.GreefF, while visiting the island of Rolas, in the Gulf of Guinea, found
some
crinoidstherewhich
heidentifiedwiththespeciesinhabit- ing southern Europe. Carpenterwas
inclined to considerthem
as beingin realitytheAntedon
diibenii ofBohlsche,butitseems probable that they are identical with the Antedon hupferi of Hartlaub,which
is closely related to that species.
DeLoriol,discussing theechinodermsofMauritiusin1883,includes, as did Michelin, Tropiometra carinata.
The work
of thetwo French
steamers, the Travailleurand
the Talisman,had
resulted in the discoveryofmany
interesting crinoids off the coasts of southernEurope and
northwestern Africa. Scat- tered references to these arefound inthe writings of E. Perrier, Cap- tain Parfait, de Folin,and
of the Marquis de Filhol, but they are mostly very indefiniteand
very unsatisfactory. Interest in these crinoids appears tohave
soon died out,and no
detailed reportupon them was
ever prepared.Prof. F. Jeffrey Bell, in listing the echinoderms obtained
by
the Alert in the western Indian Ocean, records (1884)immature
speci-mens
of a species of uActinometra^
from
theAmirante
Islands.4 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.In the
same
year P.H.
Carpenter publishedan
account of the crinoids occurringbetween
the Faroe Islandsand
Gibraltar, mostly basedupon
the results of thework
of the Porcupine expedition,and
healso finishedthemonograph on
the stalkedcrinoidswhich had
been obtainedby
the Challenger. This latter ismore
comprehensive in scope than is indicatedby
its title, for it includesan
account of the materialobtainedby
all the other exploring ships,in so farashehad
access toit,and
citesnearly allofthepublishedreferences, thusoffer- ing a reasonably completesummary
of everythingknown
in regard to the recent stalked crinoidspreviousto 1884.Four
yearslater thismonograph was
followedby
a similarwork
dealing with thecoma-
tulids; but thisis less comprehensive inits scope, so that speciesnot
among
the Challenger collections are neither described in detail nor figured.As
the Challenger obtained but a single species while in African waters, atSimons Bay,
Natal,and
that a well-knownform
(Comanthus wahlhergii), it is evident that African species receive a comparatively smallamount
of attention.Hartlaub's
monograph on
the comatulid fauna of the East Indian Archipelago, published in 1891 (after a preliminary paper in 1890),fills
many
of thegaps leftby
Carpenter in the Challengermonograph.
Two new
Africanspecies, onefrom
theRed
Sea (AntedonMunzingeri)and
the otherfrom
the Ivory coast (Antedon hupferi), are described asnew and
figured,and two
others,known from
theRed
Sea since 1817and
1833, are for thefirst time adequately figured.Bellin 1892 described a
new
species of "Antedon" from
Mauritius under thename
of A. emendatrix.Ever
since then it has been a great puzzle to determinewhat
the species really is,as the diagnosisis far
from
clear,and
thereis a possibilitythat it coverstwo
distinct species belonging totwo
different genera, oneofwhich
is a species of Cenometra,though
Bell did not recognize its affinity with C. bella,which
Hartlaubhad made known two
years previously.In 1899
Hubert Ludwig
publishedan
important paperupon
the echinoderms of Zanzibar, basedupon
a collectionmade
in that countryby
Doctor Voeltzkow. In this herecapitulates the previous records for the region about Zanzibar publishedby von
Martens,Rathbun,
Pourtales,and
Carpenter,and
addsAntedon
flagellata, a speciespreviouslyknown
onlyfrom
Singaporeand
thePelew
Islands.The work
of the steamer Valdivia while under charter to theGer-man Government
resultedin the discoveryoftwo
interestingcrinoids off the coast of Somaliland,which
are figuredby
ProfessorChun,
with tentative identificationsfurnishedby
Professor Doderlein, in his interesting account of thevoyage
(Ausden
Tiefen des Weltmeeres).One
of thesewas
a species of Rhizocrinus, a genusknown from many
places in the Atlantic, but hitherto only
known
outside that areafrom
the indefinite report of Korotneff,who
found a large speciesno. 180S.
RECENT
AFRICA'SCRINOIDS^-CLARK.
( ?R.weberi) in the Straits of
Sunda
; the otherwas
anew
species of Pentametrocrinus.The work
of the PrinceofMonaco
hasadded
a great deal to ourknowledge
of the crinoids of thedeeper watersoff northwest Africa.Much
information is contained in short papers publishedby
the Prince himselfand by
ProfessorsKcehlerand
Richard. Allthe data acquired has recently (1909) been presented in magnificentform
under the authorship ofProfessor Kcehler.Itisvery remarkablethat the Princesse-Alice should
have
obtained onlytwo new
species, but the lack of quantity ismore
thancom-
pensatedby
the interest attaching to one of them, Gephyrocrinus grimaldii, representing a second genus of Hyocrinidse, first described in a preliminary paperby
Professor Kcehlerand
Dr. F. A. Bather, jointly, in 1902.The
investigations conductedby
theCape
ofGood Hope
Govern-ment
into the marine resources of that colonyhad
resulted in the accumulationofmost
excellent collections ofechinoderms. These were turned over to Prof. F.Jeffrey Bell for determination,and
he foundamong them
four species of crinoids, three ofwhich
he described as new, assigningthem
all to systematic positions veryremote from
those they in reality occupy; his fourth specieswas
misidentified.These fourspecies are
:
Antedon
capensis=
Tropiometracarinata.Antedon
sclateri=
Pachjlometrasclateri.Antedon rnagnicirra
=
Crotalometra magnicirra.Actinometra parvicirra
=
ComantTiuswaldbergii.In his report
upon
the stalked crinoids of the Siboga expedition,which
appearedtoward
the close of 1907, Professor Doderleinfigured the species ofRMzocrinus
dredgedby
theVaUivia
off Somaliland, callingit,most
appropriately,RMzocrinus
cliuni.Mr.
&
Herbert C.
Chadwick
in 1908communicated
to the Linnsean Society ofLondon
a short paperupon
the crinoids collectedby
Mr.Cyril Crossland
on
the coast of theSudan
during the course of the investigations of the marine biology of theRed
Sea under the direc-tion of Prof.
W.
A.Herdman.
Six species are listed, four ofwhich
are
new
to the region; these are:
Antedon
serripinna=
Colobometra cliadwicki.Antedonparvicirra
=
Iridometra xgyptica.Antedon
marginata=
?Steplianometra marginata.Antedon imparipinna
=
Dichrometraprotectus.Antedon palmata
=
Dichrometrapalmata.Antedon
savignyi=
Heterometra savignii.Mr.
Chadwick
himselfwas
not certain of the correctness of his determination ofAntedon
marginata,and
in a review I took the liberty of stating thatAntedon
serripinnaand A.
parvicirra were6 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.possibly not quite the
same
as the East Indian species calledby
thosenames by
Carpenter.Thanks
to the kindnessofMr.Chadwick and
ProfessorHerdman,
Ihave
been able toexamine
one of the specimensidentified as-4. serripinna,and
I find it to be anew
species of Colobometra,and
not an Oligometra at all. I also found in the BritishMuseum
a specimen ofwhat
isundoubtedly
hisAntedon
parvicirra,
which
turned outto be anew
species of Iridometra.In October 1909, Professor Bell reported
upon
the echinoderms of thePercy
Sladen Trust expedition,which
included four crinoids.All weremisidentified.
The
fourspecies are:Actinometramultiradiata
=
Comatella maculata.Antedon
carinata=
?Cosmiometra gardineri.Antedon
palmata=
Stephanometra indica.Antedon
spicata=
Cenometra emendatrix.MATERIAL.
The
specimensexamined
in connection with the presentwork
include all the crinoids
from
Africa in the BritishMuseum,
theBergen Museum,
theMuseum
ofComparative
Zoology, the Copen-hagen Museum,
theHamburg. Museum,
theMuseum
furNaturkunde and
theMuseum
fur Meereskunde, Berlin, the OceanographicMuseum
atMonaco,
the ParisMuseum, and
the U. S. NationalMuseum. Among them
are the originals ofall the published records of Leuckart, Guerin-Meneville, J. Muller, Michelin, Dujardinand Hupe, von
Martens, WyvilleThomson,
Pourtales,Rathbun,
E. A.Smith, Bell, P.
H.
Carpenter, Hartlaub,and Ludwig, and most
of those of Doderleinand
Chadwick.At Lyons
Prof. R. Kcehlerand M. Vaney showed me
thespecimens collectedby
the Travailleur, the Talisman,and
the Princesse-Alice,upon which
they are soon to publish a report.Two
of theirnew
species Ihad
already seen in themuseums
atLondon and
Paris,and
Ihad drawn up
diagnoses ofthem;
but it is onlyfair tothosegentlemen to withholdmy
diagnoses untiltheyareable topublishtheirs,and
toconfinemyself,intreating of the material collectedby
theFrench and Monacan
ships, to the published records.FAUNAL RELATIONSHIPS OF THE AFRICAN
COASTS.There are
known
to-dayfrom
the coasts ofAfricaand
the outlying islands fifty-three species of recent crinoids; forty-five of these belong to the Comatulida, representing seven families, viz; the
ComasteridaB (six); the Himerometridre (twelve); the
Colobome-
tridre (six); theTropiometridse (three); the ThalassometridaB (eight)
;
the Antedonidse (eight),
and
the Pentametrocrinidae (two), while eight arestalked, representing the Pentacrinitidse (one); the Hyocri- nidse (one),and
the Rhizocrinidse (six).These
species are included in twenty-fourgenera, ofwhich
four arestalked.no.180S.
RECENT AFRICAN CRINOIDS— CLARK.
7'
The
African shores are faunally divisible into five well-markeddistricts, each withitscharacteristic species
and
genera, as follows:MEDITERRANEAN COAST.
Family ANTEDONIME.
Antedonmediterranea. Leptomctraplialangium.
NORTHWEST COAST(STRAITS OF GIBRALTAR TO CAPE VERDE, INCLUDING THE AZORES, MADEIRA,CANARY,ANDCAPEVERDEISLANDS).
Family COMASTERID.E.
Neocomatella
(new
species).1Family THALASSOMETRIDiE.
Crotalometraflava.1 Tlialassometra lusitanica.1 Titalassometra omissa.1
Family ANTEDONIDyE.
Antedon
bifida. Leptometra celtica.Family PENTAMETROCRINIDyE.
Pentametrocrinus atlanticus.2*
Family PENTACRINITID^.
Endoxocrinus wyvilletliomsoni}
Family HYOCRINID^E.
Gephyrocrinusgrimaldii.1
Family BOURGUETICRINIDyE.
Batliycrinusgracilis.1 Batliycrinus recuperatus.1 Bothycrinus perrieri.1 RMzocrinusparfaiti.2
1
WEST COAST (INCLUDING THE ISLANDS OF ST.HELENAANDASCENSION).
Family TROPIOMETRIDJD.
Tropiometrapicta.2
Family ANTEDONIDiE.
Antedonliupferi.
1Deep-waterspecies. 2Occursalsoin theWestIndies.
8 FROCEEDINQS OF THE NATIONAL MUSEUM.
vol.40.Family THALASSOMETRID^.
Crotalometraporrecta.1
Thalassometm
multispina.1Family RHIZOCRINID^E.
BathycrinusaldricManus-1
southeast coast (from mombasa to cape town, including all the outlying islands).
Family COMASTERID^.
Comatella maculata. Capillastermultiradiata.
Comissiaignota.
Comanthus
parvicirra.Comanthus
waMbergii.Family HIMEROMETRID.E.
Amphimetra
africana. Craspedometra madagascarensis.Ileterometra joubini. Heterometra gravieri.
Stephanometraindica. Dichrometraflagellata.
Family COLOBOMETRID.E.
Cenomctraemendatrix. Decametramobiusi.
Decametramodica. Decametra alaudse.
Oligometraserripinna.
Family TROPIOMETRILXE.
Tropiometracarinata.
Family THALASSOMETRIDyE.
Crotalometra magnicirra.1 Cosmiometragardineri.1 Pachylometrasclateri.1
Family ANTEDONIM}.
Iridometra mauritiana. Perometra afra.
NORTHEAST COAST(FROM BRITISH EAST AFRICA TO SUEZ).
Family COMASTERID^E.
Comanthus
{?parvicirra).iDeep-waterspecies.
no.1808.
RECENT AFRICAN CRINOIDS— CLARK.
Family HIMEROMETRID.E.
Craspedometraater. Diclirometraprotectus.
Heterometrasavignii. DiclirometraMunzingeri.
Stephanometramarginata. Diclirometrapalmata.
Family COLOBOMETRID.E.
ColobometrachadwicM.
Family TROPIOMETRID/E.
Tropiometraencrinus.
Family ANTEDONID^.
Jridometrasegyptica.
Family PENTAMETROCRINID^E.
Pentametrocrinus (new species).1
Family RHIZOCRINID/E.
Rhizocrinus cliuni.
Mediterranean coast.
— Sofar as we know,
the Mediterranean coast
of Africais thesame
faunally as theMediterranean coast of Europe.
Only two
genera,Antedon and
Leptometra, are found here, both ofwhich
areprimarily, as Ihave
shown, of IndianOcean
orEast Indian origin,2and
both ofwhich
occurin the Atlanticnorthward
nearly or quite toNorway.
Thesetwo
generamay
be said to delimit theEuropean
faunal area—
in reality an attenuated derivativefrom
the great Indo-Pacific-Japanese region— and this European
faunal area
may
logically be divided into (1) the Mediterranean area, inhabitedby
Antedon mediterranea, A. adriatica,and
Leptometra plialangium;and
(2) the European-Atlantic area, inhabitedby Antedon
petasus, A.bifida,and
Leptometraceltica,and
extendingfrom Norway
south toMorocco and
Madeira.Northwestcoast.
—
Thisisthemeetingground between
the precedingand
succeedinglittoral faunal areas; here the European-Atlantic area reaches its southernand
the West Africanitsnorthern limit.In the deeper water, however, inhabited
by
species of the Inter- mediate faunal zone,3we
find a stronglymarked "West
Indian"clement, indicated
by
Neocomatella, Pentametrocrinus atlanticus,and
Rhizocrinusparfaiti.1Deep-waterspecies.
2Cf.Vid.Medd.fradennaturhist. ForeningiK0benhavn,1909, p.12Set seq.
3Idem.,p. 120.
10 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol. 40.'Thanks
to thework
ofthe ships of theU.
S. CoastSurvey and
theU.
S. FishCommission we have
longhad
dataupon which
to base a concept of the Antillean crinoid fauna,and we
are able to state with a reasonable degree of accuracy that this fauna is directly derived from,and
forms the outerfringe of, the great Intermediate area, themaximum
intensityofwhich
lies within a trianglewhose
apices are,roughly,southernJapan, the
Kermadec
Islands,and
Singapore. Just as the generaAntedon and
Leptometra characterizing theEuropean
faunal area are distinct from,though
very closely related to, the parent genera Mastigometraand
Psathyrometra of theEast
Indies, so the generafound
in theWest
Indies are very closely related to,butentirely distinctfrom,thecorresponding
East
Indiangenera.The
correspondenceiswell brought outby
the followinglist:
WEST INDIAN GENUS. CORRESPONDING EAST INDIAN GENUS.
Family COMASTERIDiE.
Nemaster. Capillaster.
Neocomatella. Comatella.
Leptonemaster
.
Comissia.
Comactinia. Comatula.
Family IIIMEROMETRID.E.
Analcidometra. Stephanometra.
Family THALASSOMETRID/E.
Stylometra. Cosmiometra.
n
• . IPachi/lometra.Crinometra.
\Glyptometra.
Family ANTEDONIDiE.
Tlypalometra
.
Enjtlirometra.
Zenometra. Psathyrometra.
Coccometra. Thysanometra.
Certain other genera probably should be paired in the
same way, but
our data in regard tothem
is as yet insufficient for generaliza- tion; anumber
of genera are, of course,common
to both regions,though
the species arenever the same.Now
the entire western coastofNorth and South America
is, as Ihave
shown,1 purely Antarctic, or,more
precisely, Magellanic, in its affinities; the Indo-Pacific-Japanesespeciesdisappearfar to thewest-ward among
theSouth
Sea islands. It is thus highly improbable that theWest
Indies could everhave become
colonizedfrom
the westward; if suchhad been
the case therewould
certainly be traces1GeographicalJournal,vol. 32,No.G,p.002etseq.
no. 1808.
RECENT AFRICAN CRINOIDS— CLARK.
11 remainingofthezoogeographicprogressionon
thetropicalwestAmer-
ican coast. Moreover, the geology of the country
between North and South America shows
that there has been a fairlypermanent
land barrierat all timesbetween
theCaribbeanand
thePacific; such channels ashave
intercommunicatedbetween them
couldneverhave
been deep, asnone
of the characteristic animals occurring below the sublittoral zoneon
the Pacific side appeartohave
passed through to the Caribbean.No
crinoids ever passed through in either direction,and
there is not even a subfamilyor a genuscommon
to both coasts of Central America; but crinoids are almost entirely sessile— much
more
nearly sothanany
othermarine organisms— and many
barriers therefore operate against their dispersalwhich
are ineffective against other animals, including the otherechinoderms.1The
crinoids, therefore,move
very slowly intonew
territory for this reasonand
also because of their slight adaptability to changes of salinityand
especially of temperature,and
the fact that afew
of the littoral echinoids, ophiuroids,and
asteroids have, during the periodswhen communication was
establishedbetween
the Caribbeanand
thePacific,made
theirway from
theformerinto the latter, does not inany way
disprove the hypothesis that there has at all times been aninsuperable barrier to thedispersal of the crinoids.Thus
it seems necessary toassume
that theWest
Indian crinoid fauna reached the Caribbean basinfrom
the eastward,from
the northwest coast of Africa, having previously reached that districtfrom
thesouthward
or southeastward, notfrom
the eastward or northeastward.According to this, the northwest African intermediate fauna (including that of
Europe
south of theBay
of Biscay) is zodgeo- graphicallymore
important than theWest
Indian,though
atpresentmuch
lessknown;
the two, so far as can be judged, are practically the same, so that together theymay
be considered asmaking up
a South European-Northwest African-Antillean faunal division of the intermediate region,which
falls naturally intotwo
subdivisions, (1)an
Afro-Europeanand
(2) an Antillean.Itis avery curious fact that almost withouta singleexception the genera
and
species of thisfaunaldivision inhabitdeeper—
oftenmuch
deeper
—
water than the generaand
speciesfrom which
theyhave
been derived in the
East
Indian seas,and
theirhabitat in the eastisdeeper again than their habitat in the west. It
would
appear that this faunal division has takenfrom
the great parent Indo-Pacific- Japanese region only such genera as aremost
plasticand
can best adapt themselves to changing conditions; these generahave
been abletosurvive,buthave become
differentiatedfrom
theparentstock, while all the other less plastic genera,which we must assume
wereiVid.Medd.fradennaturhist. ForeningiK0benhavn,1909,pp.117-133.
12 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.originally carried alongwith them,
have
died out.The
difference in depth of habitatbetween
the specieson
the opposite coasts of the Atlantic is probablydue
to a difference in the nature of the food supply,and
the increased depth of the habitat of theWest
Indian genera over theirEast
Indianrelatives originated probablyfrom
thesame
cause.1There is, therefore, a close similarity in origin
between
the species oftheEuropean and
the south European-northwest African- Antillean faunal divisions; the formerhave
reached their present habitatby
passing
northwestward around
or across the northernend
ofwhat
isnow
Africa, the latterby
passingmore
to the southward.At
first similar extensionsfrom
thecommon
parent region, the diverse con- ditionswhich
inpast geologic agesthey encountered,oneinthenorthand
the other in the south, reducedthem
differently, so thatnow
they present a totally different aspect,
and
onewould
neversuspect acommon
originwere
they not both equally closely related to theparentregion.
Itisveryinteresting thatintheagessincethe constituentelements ofthese
two
faunaldivisionslefttheparentregiononly comparatively slight changeshave
occurred;no
widely diverse generic typeshave been
evolved,and none
of the collateral parent generahave
entirely died out behind them,though
theyhave
largely disappearedfrom
the intervening seasof the present day, being preserved, however, in certaincases, asfossilsin the laterrocks.West coast.
—
There are onlytwo
littoral comatulidsknown from
the west Africancoastand
the outlyingislands; oneof these (Tropi- ometra *picta) is, so far as I can see, identical with thecommonest
species
on
the opposite coast ofSouth
America, while the other (Antedonliupferi) is very close to a corresponding species originally describedfrom Rio
de Janeiroand
since reportedfrom
Abrolhosand Madeira
(Antedondiibenii).The
third littoral Braziliancomat-
ulid (Nemaster lineata) has probably spread
southward from
theWest
Indies just as Tropiometra pictahasworked
itsway northward
to thoseislands.
The
faunal division characterizedby
the presence of Tropiometra pictaand
the small shortarmed
species ofAntedon may
be convenientlyknown
as theWest African-SouthAmerican
area.In
the deep waters of the Atlantic there occur certain genera of the Oceanicregionwhich
arefound
in all deepseas, exceptthat they never intrudeupon
the territory occupiedby
the so-called Polar- Pacific species.Such
genera are Bathycrinus, Bathymetra, Orota- lometra,and
Thalassometra; possibly Gephyrocrinus should also be placed here.iGeographicalJournal,vol. 32,No.6,p.G02etseq.
no.1808.
RECENT AFRICA
XCRINOIDS— CLARK. 13
Southeast coast.— The southeastern coast of Africa is faunally the
richest section of the continent, supporting twenty-two
species dis-
tributed among
eighteen genera,of which
lattereleven do not occur
farther north. All of the genera are widely spread throughout the
Indo-Pacific-Japanese region, but sixteen of the twenty-two
species
are confined to southeasternAfrica, onlysixoccurring in Ceylonand
eastward. The
only four species of the deeper water yet known
belong to characteristicallyEast Indiangenera.
The
affinities of this district are obviously with Ceylonand
the region to the eastward of that island rather than with the coasts of theArabian
Sea,and
themodern
crinoid fauna evidentlyworked southwestward
along the linenow
indicatedby
theMaldive, Chagos,and
Cargados islands towhat
isnow
Mauritius, Madagascar,and
southeastAfrica.The
following genera arecommon
to southeast Africaand
Ceylon (or theBay
of Bengal),but do not occuron
theshores of theArabian Sea north of Ceylon ornorthof British EastAfrica:Gomatella. Decametra.
Capillaster. Oligometra.
Comissia. Crotdlometra.
BennettiagroupofComanthus. Gosmiometra.
Amphimetra.
Pachylometra.Cenometra. Perometra.
The same
is, so far aswe know,
true ofthe following species:Gomatella ma'culata. Stephanometra indica.
Capillaster multiradiata. Dichrometrajlagellata.
Oligometra serripinna.
As
the south European-northwest African-Antillean division offerssome
strikingpointsof similarity totheattenuated western extremity of the Indo-Pacific-Japanese region, as seen in southeast Africa,we
are justified in supposing that it
was
originally derivedfrom
itby
passage acrosswhat
isnow
central Africa; in other words, that the genera characterizing ithave moved outward from
the East Indian region, firstsouthwestward
to southeast Africaand
then northeast-ward
to their present habitat.The
areafrom Mombasa southward
toCape
Colony,and
including Madagascar, the Seychelles, Reunion,and
Mauritius,and
the other islands as far as theChagos
groupmay
beconvenientlyknown
asthe SoutheastAfrican faunal division of the Indo-Pacific-Japaneseregion.Along
the coasts ofCape
Colonyand
Natal occurs a comasterid {Comanthus wahlbergii) foundnowhere
else, but closely related toComanthus
trichoptera of southern Australia.Although
it is asso- ciatedwith tropical forms,and
although there areno
representatives there of the other south Australian species,we must
recognize the fact that the Cape subdivision is not quite thesame
faunally as the14 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40."shores farther north; it is poorer in species, and is modified
by
the intrusion of a south Australian element, just as has beenshown
in the case ofmany
other animals.Northeast coast.
— The northeast coast of Africais singularly differ-
entin the aspectofitscrinoid faunafrom
the coastto thesouthward.
Of
strictlycomparable
species there are only ten, distributedamong
eight genera, of
which
one is notknown from
farther south; thisis:&
Colobomctra.
All of thegenera arepurelyEast Indian; of thespeciesfiveextend to India or beyond; these are:
Heterometra savignii. Dichrometra protectus.
Stephanometra marginata. Dichrometrapalmata.
Tropiometraencrinus.
The
fauna ofnortheast Africa thereforeis purely a derivativefrom East
Indian stock, just as that of southeast Africa is, but a curious segregation of the generaand
speciescomposing
the Indo-Pacijic- Japanese region has occurred, one set of forms following the coasts of theArabian
Sea, the other extending in a southeasterly directiontoward
theCape.While
thefaunasofthenortheastand
ofthe south- east coasts differ considerablybetween
themselves, they are both aboutequally related to the generalEast
Indian fauna,and
thecom-
ponent species of both are remarkable in being in general smaller than the corresponding species in the great parent area.The
shoresfrom
Somalilandnorthward
toand
including theR6d
Sea,
and
thence eastward to the Persian Gulf,may
be considered asmarking
the NortheastAfrican faunal division.Whether
or not the generaAntedon and
Leptometra reached their present habitatby way
of theArabian Sea—
that is, through the faunanow
occupying the shores of thatbasin—
it is not possible to say; Mastigometraand
Psathyrometra, theireastern equivalents, occur at or near Ceylon, but areknown no
farther west.They may have
gone"overland" from
India;may have
passed along the shores of theArabian
Sea (or its ancienthomologue) and
subsequently died out,or,which
ismuch
themost
likely, theymay
yetremaintobedis-covered
between
Ceylonand
Suez.Summary.— The
crinoid faunaof the coastsof Africafalls into five divisions, as follows:1.
The European
division,including the Mediterraneancoast,and
thenorthwestcoastnorthofMorocco and
Madeira; thisfallsintotwo
subdivisions:(a)
The
Mediterranean subdivision, comprising the coast of the Mediterranean,and
(b)
The
European-Atlantic subdivision, extendingon
the Atlantic coastfrom Madeira and Morocco
northward;no.1S0S.
RECENT AFRICAN CRINOIDH—
VI.M{
Ik. 152.
The South European-Northwest African-Antillean
di- vision, extendingfrom Madeira and Morocco northward
to theBay
of Biscayand westward
toand
including the Caribbean Sea;3.
The West African South-American
division, including the coast south ofMorocco and
the opposite coast of Brazil, with the interveningislands;4.
The Southeast African
division, extendingfrom Mombasa
toCape Town, and
eastward to include Madagascar, the Seychelles, Reunion,and
Mauritius,and
the interveningislands eastward to theChagos
archipelago; thesouthern partof this division,comprising the coasts ofCape
Colonyand
Natal, forms the Capesubdivision;and
5.
The Northeast African
division, extendingfrom
Somalilandnorthward
throughout theRed
Seaand
eastward to thePersianGulf.The West
African-SouthAmerican
division is an attenuated off- shootfrom
theSouth European-Northwest
African-Antillean divi- sion,which
itself is derivedfrom
the Southeast African division, the last thesomewhat
modified southwestern extremityofthe greatIndo- Pacific-Japanese faunal area.The European
faunal division is probably an attenuated offshootfrom
the Northeast African division,which
itself is the considerably modified northwestern extremity of the great Indo-Pacific-Japanese faunal area.ANNOTATED
LISTOF
SPECIES.I.
COMATULIDS.
SuborderOUGOPHREATA.
Family COMASTERIDJE.
Subfamily CAPILLASTERINJE.
Genus NEOCOMATELLA
A.H.
Clark.NEOCOMATELLA(newspecies).
Actinometrapulchella (part) P. H. Carpenter, Proc. Roy. Soc. Edinburgh,vol.
12, 1884, p. 369; Challenger Reports,vol. 26,Zoology, 1888, p.304—Kcehler andVaney, Bull,dunius. d'hist. nat., 1910, No.1, p. 32.
Localities.—-West of Gibraltar (lat. 35° 56' N.; long. 7° 06' W.);
northeast of the Seine
Bank
(lat. 34° 57' N.; long. 11° 57'W.): offCape
Penas; south ofCape
St. Vincent (lat. 35° 26' N.; long. 9° 09' W.); south oftheCanary
Islands (lat. 25° 41' N.; long. 18° 16' W.).Depth.—228-533
fathoms.Bottom temperature.
—
50.5° Fahr. (one record).Bottom.
—
Clay (onerecord).1iComatulasolariohas beenrecordedfromthe east coastofAfrica (Zanzibar)byProfessorvon Martens, butthe recordappearstohave been basedupona specimenofTropiomctracarinata.
16 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.Genus COMATELLA
A.H.
Clark.COMATELLA
MACULATA
(P.H.Carpenter).Actinometra multiradiataBell, Trans. Linn. Soc.(Zool.), (2),vol. 13, 1909, pt.
1, p. 20.
Localities.
—
Salomon,from
the reef; Coin Peros.Remarks.
—
-The specimenfrom Salomon, which
Iexamined
at the BritishMuseum,
has twenty-sixarms
85mm.
long; the cirri areXXI,
15-18.The
IIIBr series are all developed externally.Com-
pared directlywith the typeof Carpenter's maculata, thisspecimen isfound to differonly in having
more numerous
arms.Genus CAPILLASTER
A.H.
Clark.CAPILLASTERMULTIRADIATA(Linnaeus).
Asterias pectinata (part) Linnaeus, Syst. Nat.,10th ed., 1758, p. 6G3(reference to Petiver).
Asterias multiradiata Linnaeus, Syst. Nat., 10th ed., 1758, p. 663 (type-speci-
menatLund, butnot referencescited).
Comatula fimbriata Lamarck, Hist. nat. des animaux sans vertebres, vol. 2, 1816, p. 535.
Comatulacoccodistoma(ParisMuseumMS.)Dujardin and Htjpe, Hist. nat. des zoophytes; eohinoderrnes, 1862, p. 208.
Actinometra coppingeri Bell, Proc. Zool. Soc. London, 1882, p. 535.
—
Rep.Zool.Coll. H.M. S. Alert, 1884, p. 168, pi. 16,fig.B.
Actinometra multiradiata P. H. Carpenter, Challenger Reports, vol. 26, 1888, Zoology, p. 322,pi. 66, figs. 1-3.
Capillastcr multiradiataA. H. Clark, Vid. Medd. fra den naturhist. Forening i
Kjzfbenhavn,1909, p. 134.
Localities.
— Madagascar; Cape St. Andre, Madagascar.
Depth.
—
Littoral,and down
to about 30 meters.Inthe Paris
Museum
there arethree specimensof this speciesfrom
Madagascar; one has the cirriXV,
21-23,and
nineteen arms, one IIBrseries being lacking;two
of theIIBr
series are 2, the remaining seven being 4(3+4);
another (Cape St.Andre;
about 30meters) has the cirriXVIII,
21-22,and
twelve arms, one ray bearingtwo
IIBr4 (3+
4) series; the brachialsareveryshortand
overlap rather strongly;
the third islike the first,
and
also has nineteenarms
; no IIIBr series arepresent.Compared
directly with a typicalexample from
the Straits of Sunda,these threespecimensareseentobesmallerand
proportionately weaker, but otherwise I can findno
differences whatever. IIIBrseries, very
common among
East Indian specimens,do
not occur.Very
possibly these specimens represent a local permanently small race confined to the southeast African region. If this should prove tobe
the case thename
coccodistomawould
be applicable to it.no. 1808.
RECENT AFRICAN CRINOIDS— CLARK. 17 Genus COMISSIA
A.H.
Clark.COMMISSIAIGNOTA,newspecies.
ActinometraBell, Rep. Zool. Coll.II.M. S. Alert, 1884, p. 510.
Actinometrapectinata Brit. Mus.MS.
Description.
—
Centrodorsaldiscoidal,withaflatdorsalpole1.5min.- 2mm.
in diameter.Cirri
XII-XV,
9-11 (usually 10-11), 4mm.-5 mm.
long; firstsegment
short,second notquite solong as broad, the followingincreas- ing inlength tothe fourth,which
is about three times as long as themedian
diameter; the sixthand
following are about aslongasbroad;the third
and
fourth are strongly "dice-box shaped;" the fifthand
followingeach bear aminute
sharpsubterminialtubercle; the shorter distalsegmentsaresomewhat
compressed laterally,and
hence appear comparatively broad in lateralview; the opposingspine isslightand
inconspicuous.Radials concealed;
IBr
x well separated, the interradial angle of separation being about 90°.
Ten arms
30mm.-35 mm.
long, resembling in general those of Leptonemastervenustus, but proportionately slightly stouter; genital glandsaredevelopedon
the pinnules.Locality.
—
Marie LouiseIsland,and
Isledes Neufs,Amirante
group;
collected
by H. M.
S. Alert.Depth.
—
17 fathoms.Bottom.
—
Coral.Subfamily COMASTBRIN.S].
Genus COMANTHUS
A.H.
Clark.Subgenus
COMANTHUS.
Specificgroup
BENNETTIA.
COMANTHUS
(BENNETTIA)WAHLBERGII(J.MUHer).Comatulacoccodistoma(part)ParisMuseumMS.
Alecto wahlbergiiJ.Muller, ArchivfiirNaturgesch.,1843,Heft1,p. 131.
Actinometra parvicirra (part) P. H. Carpenter, Challenger Reports, vol. 26, Zoology, 1888, p.338.—Bell, Marine Investigationsin SouthAfrica, vol.4, pt.4,p. 141.
Localities.
—
Port Natal;Simons
Bay, Natal; FalseBay; Cape
ofGood Hope; Algoa
Bay, close toRiy Bank
(lat. 33° 58' S.; long.25° 51'
30"
E.); Tugela Rivermouth
bearingNW. by
W., 3£ miles distant.Depth.
—
Littoral,and down
to25 fathoms.Bottom.
— Dark sand, black specks, and
rocks; rocks.
Remarks.
—
Hitherto this species has always been confused with thecommon
East IndianComanthus
parvicirra,from
which, however,it is very sharply separated, not even belonging inthe
same
specific group. Itsclosest affinity iswithComanthus
trichoptera ofsouthern80796°—Proc.N.M.vol.40—11 2
18 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.Australia,
and
itspresence at theCape
indicates aconnectionbetween
the crinoid faunas of thesetwo
localities similar to thatshown
inmany
other groups.Comanthus
wahTbergii is a small stout species with well-developed,permanent
cirri; these arefrom XII-XXV,
13-17 (usually aboutXV,
16), about 10mm.
long;from
the fifth to the seventh (usually the sixth) a transitionsegment
is developedwhich
is usually wellmarked.
The
centrodorsal is thin discoidal, with a broad flat dorsal pole 4mm.
in diameter, resembling that of C. trichoptera.The arms
arefrom
thirteen to twenty-one innumber
(usuallyfrom
sixteen to twenty)and
arecommonly
about 60mm.
long.IIIBrseriesare rare. Allthe divisionseriesare 4 (3
+
4).The arms
areshortand
stout,taperingrapidly,much
as inComatulabracJiiolata.The
brachials overlap conspicuously,and
the ends ofthe elements of the division series are prominent.The
division series are broadand
are close together laterally.One
of the specimenswhich
Iexamined
in the BritishMuseum
(catalogued
under
thename
of Actinometra paucicirra) has thirteen pentacrinoids attached to the cirri; these are similar to those ofComactiniameridionalis
and
possess seventeen columnars.Specificgroup
VALID
IA.COMANTHUS
(VALIDIA)PARVICIRRA(J.Miiller).AlectoparvicirraJ. Muller, Archiv fur Naturgesch., 1841, Heft1, p. 145.
Alecto timorensisJ. Muller, Archiv fur Naturgesch., 1841, Heft1, p. 145.
Comatula brevicirra Dujardin and Hupe, Hist. nat. des zoophytes; echino- dermes, 1862, p. 208.
Comatulasimplex Dujardinand Hupe, Hist. nat. des zoophytes; echinodermes, 1862, p. 208.
Actinometra trachygaster (part)Lutken,Mus. Godeffr.Cat., vol.4,1869, p. 125.
Actinometraintricata (part) Lutken, Mus. Godeffr.Cat., vol. 5, 1874, p. 190.
ComatulamertensiGrube,J.B. derschls.Gesellsch.fur vaterl.Cultur,1875, p.74.
Actinometra armata W. B. Carpenter, Proc. Roy. Soc. London, vol. 24, 1876, p. 451.
Actinometrapolymorpha P. H. Carpenter, Journ. Linn. Soc. (Zool.), vol. 13, 1877, p. 440.
Actinometra meyeri P. H. Carpenter, Journ. Linn. Soc. (Zool.), vol. 16, 1882, p. 525.
Antedonmertensi Bell, Proc. Zool. Soc. London, 1882, p. 535.
Actinometra mutabilisvonGraff, ChallengerReports,vol.10,Zoology,1884, p.13.
Actinometra annotea Bell, Sci. Trans.Roy.DublinSoc. (2),vol.3, 1887, p. 645.
Actinometra elongata P. H. Carpenter, Challenger Reports, vol. 26, Zoology, 1888, p. 311.
Actinometra simplex P. H. Carpenter, Challenger Reports, vol. 26, Zoology, 1888, p. 312.
Actinometra quadrata P. H. Carpenter, Challenger Reports, vol. 26, Zoology, 1888, p. 331.
no.1808.
RECENT AFRICAN CR1NOID8— CLARK. 19
Actinometra parvicina (part) P. H. Carpenter, Challenger Eeports, vol. 26, Zoology, 1888, p. 338.
Actinometra rotalaria P. H. Carpenter, Challenger Reports?, vol. 26, Zoology, 1888, p. 313.
Actinometra guttata (Lutken MS.) Hartlaub, Nova Acta Acad. German., vol.58, No. 1, 1891, p.96.
Comatulaorientalis A.II. Clark, Proc.U. S. Nat. Mus., vol. 33, 1907, p. 155.
ComatulahelianthusA.H. Clark, Proc.U. S. Nat. Mus.,vol. 34, 1908, p. 440.
ComanthusrotalariaA. H. Clark, Smiths. Misc. Coll. (QuarterlyIssue), vol. 52, pt.2, 1908, p. 205.
Comanthus (Comanthus) rotalaria A. H. Clark, Vid. Medd. fra den naturhist.
Foreningi K0benkavn, 1909, p. 144.
Locality.
—
Seychelles;two
specimens in the BritishMuseum
were collectedby
theSea Lark
expedition,under
Prof.J.StanleyGardiner.Depth.
—
34 fathoms.Remarks.
— Inthe BerlinMuseum
there aresome immature
comas-
terids
from
the Seychellesand some
othersfrom
theRed
Sea, the latter collectedby
Dr. R.Hartmeyer, which may
belong to this species.Accordingto the data given
by
Carpenterinthe Challengerreport, the first availablename
for this species is Comatula rotalaria ofLamarck,
Carpenter's figure of Actinometra rotalaria clearly repre- sentingthis species.Upon examining Lamarck's
types at the ParisMuseum,
however, Ifound
that his rotalaria is the species called Actinometrajukesiiby
Carpenterand
Actinometrapaucicirraby
Bell, so that Miiller's parvicirrabecomes
the earliest availablename
for the present form.Miiller's Alecto wahlbergii
must
be eliminatedfrom
thesynonymy
of
Comanthus
parvicirra, inwhich
itwas
includedby
Carpenter, as it hasproved
to be quite a different thing, a species related to C.trichoptera of southernAustralia
and
not to C. parvicirra at all.COMANTHUS
(?species).Locality.
— Cape St. Andre, Madagascar.
Depth.
— About 30 meters.
Remarks.
—
Dr. P.R. Joly dredged in 1901 asinglesmall specimen,undergoing adolescent autotomy, of
some
species of Comanthus.The
cirri are XIII, 13-14; there are twenty-three arms; one of the internal IIIBrseries is 2, all the other division series being 4 (3+
4).The
raysbear 4 (3+
1), 5 (1+
4),5 (4+1),2,and
7 (4+
3)arms
respec- tively.The
cirri are rathermore
developed than is usual in C.parvicirra, with
which
thespecimen otherwise agreesfairly well,and
are rathermore
compressedand
curved distally.The
interradial perisome isstrongly plated.20 PROCEEDINGS OF THE NATIONAL MUSEUM.
vol.40.COMANTHUS
(? species).Actinomelra cumingii Brit. Mus.MS.
Locality.
—
Mauritius.Remarks.
—
There aretwo
small comasteridsin the BritishMuseum from
Mauritius, one with thirteen, the other with fourteen arms,which
are very close to, if not thesame
thing as,Comanthus
parvicirra.
Family HIMEROMETMD^.
Subfamily HIMEROMBTRIN^.
Genus AMPHIMETRA
A.H.
Clark.AMPHIMETRA
AFRICANA, new species.Description.
—
Centrodorsal, thin-discoidal, broad, the flat polar area being 4mm.
in diameter; cirrus sockets in a singlemore
or less irregular closelycrowded
marginal row.Cirri
XXIII,
29-32 (usually 30),20mm.
to 23mm.
long; firstseg-ment
short, the following gradually increasinginlengthtotheseventh or eighth,which
isaboutas longasbroad; followingsimilar, after the eleventh or twelfth very gradually decreasing in length so that the terminal segments are about one-third broader than long; tenth or eleventh (usually the latter)and
following bearing longand
promi- nent dorsal spineswhich
begin abruptly; the cirrido
not taper distally.Radialsentirelyconcealed;
IB^
very short,bandlike,in apposition laterally;IBr
2 short,two and
one-half times as broad as long, the lateral edges not so long as those of theIBr
x, in apposition; IIBr 4 (3+
4) (eight IIBr series are present; the otherarms
are broken off beforethe first syzygy); IBr, IIBr,and
first brachials in close lateral appositionand
sharply flattened againsteach other;sides ofIBr and
IIBr series slightly produced; synarthrial tubercles obsolete.Nineteen (?twenty)
arms
(in the type) apparently about 125mm.
long; first brachial short, slightly
wedge
shaped, twice as broad as longexteriorly, interiorly united; second brachial slightly largerand more
obliquelywedge
shaped; thirdand
fourth (syzygial pair) slightly longer interiorly than exteriorly, twice as broad as long interiorly; following four orfivebrachialsoblong, threetimesasbroad as long, thenbecoming
obliquelywedge
shaped, twice as broad as long,and
graduallyless obliquelywedge
shaped, after theend
ofthe proximal half of thearm
being practically oblongand
very short.Pj12.5
mm.
long, slender, flagellate distally, with twenty-nineseg- ments, at first twice as broad as long,becoming
about as long as broadon
the eighthand
nearly twice as long as broad terminally;second