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Full text of "Biologia Centrali-Americana; [or, Contributions to the knowledge of the fauna and flora of Mexico and Central America]"

LIBRARY 

UNIVERSITY OF CALIFORNIA 

DAVTS 



Digitized by the Internet Archive 

in 2007 with funding from 

IVIicrosoft Corporation 



http://www.archive.org/details/biologiacentraliOOgodmrich 



B I O L GI A 

CENTRALI-AMERICANA. 



ZOOLOGY, BOTANY, 

AND 

ARCH/EOLOGY. 

EDITED BY 

FREDERICK DUCANE GODMAN, D.C.L., F.R.S., 

AND 

OSBERT SALVIN, MA, F.R.S. 



INTRODUCTORY VOLUME. 

BY 

FREDERICK DUCANE GODMAN, D.C.L., F.R.S. 



1915. 



LIBRART 

UNIVERSITY OF CALIFORNIA 
DAVig 



II 



To My Beloved Wife 

ALICE MARY GODMAN 

who has taken the deepest interest 

and given me 

much assistance and sympathy 

in the completion of 

this work 



ai 



B 936 



CONTENTS. 



Page 
Prkface vii 

Plates I., II. 

Introduction 1 

Physical Features, etc., of the Area treated 13 

Sources from whence our Material was obtained 44 

Itinerary of Mr. G. C. Champion's Travels in Central America, 1879-1883 46 

List of Completed Volumes, Zoology, Botany, and Archaeology 5.5 

Analysis of Contents of each Volume 56 

Summary of Contents of each Volume 85 

Origin, etc., of the Fauna and Flora of Central America : — 

Fauna : 

Mammalia, By R. I. Pocock 87, 142 

Eeptilia, Batrachia, and Pisces. By C. Tate Regan 105 

Arachnida (Opiliones and Acari excepted). By R. I. Pocock 118, 143 

Chilopoda, Diplopoda, and Prototracheata. By R. I. Pocock 133 

Flora. By W. B. Hemsley 145 

Maps I.-VIII. 



CORRIGENDUM. 

Page 55, line 11 from top. For Protobracheata read Prototraclieata. 



PREFACE. 



I PEEL that an apology, as well as an explanation, is due to the subscribers of the 
' Biologia Centrali-Americana ' for the length of time that has been occupied in 
the production of this work. When it was commenced, in September 1879, it was 
estimated in our prospectus that, when completed, the Zoology would "not much 
exceed sixty parts, equivalent to about twelve volumes of 500 pages each," and that 
twenty parts would suffice for the Botany, the two subjects to be issued concurrently. 
The inclusion of the Archaeology was not at that time contemplated, and this subject 
was only undertaken later in consequence of the investigations made by my friend 
Mr. A. P. Maudslay of the famous ruins in Central America, which, together with his 
beautiful photographs, made a valuable addition to our knowledge of the country. 
We had, moreover, underestimated the vast amount of additional material which 
subsequently came into our possession, and thus necessitated the extension of the 
Avork to a total of 215 parts, or 63 volumes. 

It had been our intention at the termination of the work, and after a careful study 
of the Zoological and Botanical material accumulated from this hitherto little-known 
but exceedingly rich country, to have summarised tlie result and discussed its bearing 
on the interesting subject of geographical distribution. Salvin's death after a long 
illness, and my own advancing years and ill-health, compelled me to abandon this 
project, and I should have been obliged to content myself with the conclusions arrived 
at by the various contributors in their respective Introductions had it not been for the 
assistance of Messrs. R. I. Pocock and C. Tate Kegan, to whom I am greatly indebted for 
their respective articles on the Mammals, Reptiles, Fishes, Arachnida, Chilopoda, etc., 
which are included in the Indroductory Volume. As regards the Insecta generally, 
which occupy such a large portion of the work, so little is as yet known of the fauna 
of other tropical regions that no satisfactory comparison can be made. 



viii PEEFACE. 

Mr. W. B. Hemsley, who had previously contributed the volumes on the Botany, had 
almost completed an article on the geographical distribution of the Flora, practically 
bringing this subject up to date, when, I regret to state, his health completely failed, 
and he was reluctantly obliged to relinquish his task. Recently he has, however, 
been able to furnish me with a precis of his conclusions, which forms a valuable 
addition to our knowledge of the subject. 

It now only remains for me to offer my grateful acknowledgments to all those 
who have assisted me with their various contributions, and without whom the work 
could not possibly have been undertaken. To my Secretary, Mr. G. C. Champion, 
I am specially indebted for the valuable assistance he has rendered as collector, 
contributor, and also as subeditor, in which last capacity his advice has been of 
inestimable value. His knowledge of Entomology, especially of Coleoptera and 
llhynchota, has made him one of our most important contributors, and he has 
either undertaken alone, or shared in the production of, no less than nine volumes 
of the ' Biologia,' My warm thanks are also due to my assistant, Mr. A. Cant, who 
has given very important help during the progress of the work, both in setting the 
insects, in labelling and arranging them, as well as in making very careful dissections 
and preparing slides for microscopic examination. 

F. D. G. 

June 1915. 



PLATE I. 




f.0.(v^ 



(KM 






PLATE II. 



^ 





I 



INTRODUCTION. 



A siiOKT account of the events that led to the publication of the ' Biolo^ia Centrali- 
Amcricana ' may be of interest to our readers, and I will therefore first give a sketch 
of the early days of Salvin and myself, so far as they have a definite bearing on the 
study of Natural History, and of the circumstances which drew our attention especially 
to Tropical America. 

Osbert, the second son of Mr. Anthony Salvin, the eminent architect, was born at 
Finchley in 1835 and educated at Westminster and Cambridge. That he developed 
a very early taste for natural history is clear from the series of bird skins, now in 
the Natural History Museum, collected by him as a boy and labelled ' Finchley.' 

I, Frederick DuCane Godman, third son of Joseph Godman, of Park Hatch, Surrey, 
was born in January 1834, and at the age of ten went to Eton, but three years later 
a very severe attack of what was then called low fever necessitated my removal, 
and for some years I was unable to work at all. When my health was sufficiently 
re-established, I received instruction from tutors until I was eighteen years old, when 
1 made a trip to the Mediterranean and Black Sea, visiting Gibraltar, Southern Spain, 
Athens, and Constantinople en route. During the time spent at home I interested 
myself in Natural Plistory, paying special attention to the British Mosses and Ferns, 
of which I made a considerable collection. Birds were always a source of deliglit 
to me, and I could recognise a large number of British species as well by their flight 
as by their note. 

In 1853 I entered Trinity College, Cambridge, as an undergraduate, and Salvin, in 
the following year, went to Trinity Hall, of which College he became a scholar ; he 
graduated as a Senior Optime in the Mathematical Tripos and was afterwards made 
an Honorary Fellow. With similar tastes, it was only natural that we soon met 
and became fast friends, thus forming that close intimacy which only terminated 
with Salvin's death on June 1st, 1898. Salvin was a skilful mechanic, and very 
ingenious in carpentry and cabinet making. Whilst still at Westminster, with the 
assistance of his elder brother, he built a boat thirty feet long and fitted it with 
a steam engine, the wliole of which, with the exception of the boiler, was made 
by the two brothers. This boat was launched on the river, and in it they went to 
a Thames regatta, but, having tested its powers and proved its capability, their object 

BIOL. CENTR.-AiiER., Introd. Vol., January 1915. B 



-i lA'TEODUCTION. 

was achieved, and the boat was finally sold. Some years afterwards, at Diicnas, in 
Guatemala, when Ave required specimens of the duck and waterfowl which frequented 
the neighbouring lake, Salvin again turned his hand to boat building. Tiiis time the 
ribs and frame were made of sticks of green wood cut and fastened together ; over 
this, the hair having previously been removed, a raw ox-hide was drawn, and as the 
hide shrank, it bound the whole tightly togethi r and made an excellent boat, easily 
accommodating two pe-iple. In this craft we had many sails upon the lake and 
obtained examples of the birds resorting there. 

During our College da}s, Salvin and I made frequent expeditions together to the 
fens and other places in the neighbourhood of Cambridge, in order to collect birds' 
eggs and lepidoptera. On one occasion we heard of a bustard which had been seen 
in Wicken Fen, and we spent a couple of days searching for it, but with no intention 
of shooting so rare a visitor. We found both its foot-tracks and some shed feathers, 
but, as we learned afterwards, the bird had been shot at and probably wounded by 
one of the fenmen, as it was never seen again. We also spent our leisure hours in 
Baker's shop, the well known bird stuffer in the Trumpington Road, skinning and 
setting up birds — an experience which we found of great service to us afterwards when 
in Central America and on other expeditions. 

A good deal of fenland being then undrained, Swallow-tailed butterflies were always 
to be found, and we collected the larvae and bred them in Baker's shop. The 
' Large Copper' had so recently become extinct, that we searched in vain for it, though 
Brown, the tailor in Cambridge, who was an ardent British lepidopterist, had a long 
series in his cabinet, mostly specimens bred from the larvae he had collected a few 
years previously. 

While still at Cambridge there were several other University men keenly interested 
in Ornithology, notably the two brothers Newton, Simpson, and my brother Percy, and 
after our spring rambles we used to meet in each other's rooms and discuss the result 
of our various expeditions. It was at one of these meetings in 1857 that it was first 
suggested that some record should be kept of these proceedings, and the idea of 
establishing a Magazine solely devoted to Ornithology was mooted, but nothing 
further was done till November 17th in the following year, when a meeting took 
place in Alfred Newton's rooms in Magdalene College, at which Salvin and myself, 
Simpson, Wolley, Sclater, Newton, and other ornithologists were present. Before 
the party broke up it was resolved: "That an Ornithological Union of twenty 
members should be formed, with the object of establishing a new Journal devoted to 
Birds : that Lieut.-Colonel H. M. IJrummond should be President, Professor Newton 
the Secretary of the Union, and P. L. Sclater should edit the Journal : that the title 
of the Journal should be ' The Ibis.' " 

The first volume of ' The Ibis ' appeared in 1859, and the Magazine has now reached 
its 56th volume, and the Union has over four hundred and forty members. 



I^'T]{ODUCTIOX 8 

In 1857, Salvin made a birds'-nesting expedition witli the Rev. H. B. Tristram and 
Mr. W. H. Simpson (afterwards Huddleston) through Tunisia and Eastern Algeria, in 
which I was to have joined them, but an accident in the hunting-field laid me up 
for some weeks and prevented me from accompanying them. The result of this five 
months' journey forms the subject of two valuable papers, one by Salvin, the other by 
Tristram, publislied in the first volume of 'The Ibis.' liater in the year, when I liad 
sufficiently recovered from my accident, I went with my brother Percy to Bodo, in 
the north of Norway ; there we remained for some weeks exploring the surrounding 
country and were fortunate enough to meet with and secure the eggs of the Great 
Snipe {Scolopax gallinago). Taking the steamer northward to the Alten River, we 
crossed Lapland on foot to Hapaianda, on the Gulf of Bothnia, paying John Wolley a 
short visit at Muonioniska. Before returning home we visited Stockholm, St. Peters- 
burg, and Nijnei-Novgorod. A short paper on the birds obtained on this journey 
appeared in 'The Ibis' for 1861. 

In the autumn of 1857 Salvin paid his first visit to Central America, in company 
with !Mr. George Ure Skinner, a gentleman well known to both Botanists and 
Ornithologists through the collections of rrchids and birds he had brought from tliat 
country on previous expeditions. Salvin undertook the journey, at the request of 
Messrs. Price & Co., to examine and report upon the nuts of a palm which it was 
thouglit miglit be used in the manufacture of candles. The ])alm-nuts, however, 
proved to be useless for practical purposes, and Salvin spent the remainder of his time 
in travelling tlirough the country and making a collection of birds and insects. He 
reached Belize, British Honduras, in December 1857, and after spending a few days 
there, i)roceeded down the coast to Yzabal and thence by easy stages to Guatemala City, 
making Uueuas, 30 miles south-west of the capital, his headquarters for six months. 
Salvin made two excursions to the Pacific coast region and one to the Lake of Atitlau 
in the 'Altos.' Leaving the country towards the end of June 1858, he returned to 
England via San Jose and Panama. On his return he published a paper in ' The Ibis,' 
in conjunction with Mr. P. L. Sclater, on the Ornitholo;iy of Central America (not 
including ISIexico), in which the authors enumerated 381 birds, all that were then 
known to inhabit that country. 

What he saw, however, on this expedition so whetted his appetite that he returned 
again to Guatemala in the spring of 1859, with the sole object of studying Natural 
History. He revisited Duenas, and collected in the neighbourhood for some months. 
In October he went to San Geroninio, Cohan, and other places in Vera Paz, returning 
to Duenas about the end of the year. In March I860, he was again in Alta Vera 
Paz, at Cohan, Lanquin, &c., and left for home, via Belize, in the following month. 
On this occasion he added very considerably to his collection of birds, as well as 
insects, and as a result wrote various papers in 'The Ibis' fvir 1860 on his 
discoveries. 

b2 



4 INTRODUCTIOX. 

In August 1861, I joined Salvin on his third expedition to Guatemahi, and, after 
spending three weeks in Jamaica en route, we landed at Belize ; thence taking our 
passage in a coasting schooner we arrived at Yzabal on the Golfo Dolce. Here we 
remained a few days, making preparations for our journey and engaging Indians and 
mules to transport ourselves and our baggage to the interior. This place will, 
however, always be associated in my mind with my first sight of a living example 
of one of the most striking and gorgeous of all butterflies, Morpho feleides. I was 
sitting on the trunk of a fallen tree in the forest, when it came floating past me, but 
I was so overcome with astonishment and delight at this wonderful vision that, 
although I had a butterfly net in my hand, I was utterly unable to rise in pursuit 
until it was too late to capture it. 

Crossing the Mico range of mountains, we spent a few days at Quirigua, where 
I first encountered the great Howling Monkey {Mycetes villosus), whicli frequents 
the dense forest in troops, making night hideous with its howls, which could 
be heard in the stillness for a distance of some miles. As we were sleeping in 
hammocks swung from the boughs of trees, we were somewhat disturbed in our 
slumbers. We also spent our time photographing the old Indian ruins and exploring 
the forest in the vicinity. Dry and highly sensitive plates, such as are in ordinary 
use now, did not then exist, and every photographer was obliged to carry about tlie 
necessary materials for preparing and developing his own plates, which might either 
be wet, entailing immediate development, or dry (tanning), when they could be kept 
for some days. The whole, photographic apparatus, including chemicals, fitted into 
a case, which formed a load for one of our Indian carriers. Our first essay at photo- 
graphy in the forest was not a success, as we found tliat after exposing the plates 
for twenty minutes no details were to be seen. 'I'his we discovered was owing to 
the dense green foliage overhead, through which the light had scarcely any eff"ect iu 
dissolving the nitrate of silver on the plate, and consequently no image was produced. 
In order to overcome this difficulty, we then hired Indians to cut down the trees which 
shaded the objects we wished to photograph. This delayed us a few days, which, 
however, we employed in collecting birds and insects, until a sufficient number of 
trees were felled to admit light upon the ruins, when we again proceeded to take 
photographs of the large monoliths, now obtaining very successful results. These 
ruins are fully described and illustrated in the ' Archajology ' of the ' Biologia,' by 
A. P. Maudslay. 

From Quirigua we again took the mule track, for it could hardly be called a road, 
through the valley o*f the Motagua River to Zacapa, and thence to Guatemala City. 
We spent a day or two at the Capital and then proceeded to Duenas, where we 
remained for some weeks in most delightful quarters at the house of Mr. William 
Wyld, a friend of Salvin's, Our time at this place was devoted to collecting, chiefly 
in the high forests of the Volcan de Fuego, the peak of which we ascended, and 



INTEOUUCTION. 5 

I made a separate expedition to Escuintla in the Pacific Coast re<j;ion. After our stay 
at Duenas we retraced our steps to the Capital, and, crossing the Chuacus Range into 
the plain of Salama, we took up our abode at the Hacienda of San Geronimo. Here 
we resided for some weeks, finding several species of birds and insects which we had not 
previously obtained. From San Geronimo we went to Coban, and, after spending some 
time collecting in the neiglibourhood, Ave visited Cubilguitz and Choctum in the low 
damp forest of Alta Vera Paz. At Cubilguitz, unfortunately, I contracted a sharp 
attack of fever, which obliged me to remain for some days at Coban to recruit and 
prevented my accompanying Salvin on his long and arduous journey on foot to Peten. 

When I had recovered sufficiently I returned to San Geronimo and then went to 
Buenaventura on the upper waters of the Motagua River, there called the Rio Gi-ande, 
where I employed Indians to poison some nine miles of the water in order to make a 
collection of the fish. Before commencing operations I noticed one of the ' mozos ' 
lying flat beside the river, wafting some burning material over the surface of the water, 
and, upon questioning him, I elicited that he was propitiating the spirit of the river 
in order that success might attend his efi'orts and the fish be permitted to die. The 
method adopted for this purpose was to beat the plant {I'ephrosia toxicaria, Pers.*, 
B. C.-Am., Bot. i. p. 258) on the rocks until a froth not unlike soap-suds was formed, 
this when mixed with the water caused the fish to sicken and come to the 
surface. At intervals V-shaped wicker guides were placed, so that the fish floated 
down to the point of the V, where they were then collected in baskets, and when not 
otherwise required were used as food by the Indians, who considered them perfectly 
wholesome, ^'ast numbers were thus obtained, and from them I made a selection and 
preserved a good many specimens in spirit (aguardiente), but was somewhat disappointed 
to find there were but few species represented. On my return to the Capital I journeyed 
to the Alotepeque silver-mines in company with the manager, and thence to Copan, 
Honduras, where, after spending a couple of days in examining the interesting ruins, I 
proceeded via Zacapa to Yzabal, and there met Salvin on his way back from Belize. 
Here we again parted, 1 returning to England, while he started for Duefias and the 
interior, passing through the ' Altos,' staying at Totonicapam and Quezaltenango, 
and making expeditions to the Costa Grande, Retalhuleu, and the lagoons of the 
Pacific coast at Huamuchal, close to the Mexican frontier. Salvin returned home 
early in 18G3. 

During our outward journey to Guatemala in August 1861 Salvin and I passed 
through the Azorcan Archijjelago, and I then wished I could stop and explore these 
islands, but onward we went and soon again lost sight of land. I frequently 
throughout the voyage recalled these isolated islands and determined, should 
opportunity occur, that I would explore their fauna and floia at some future date. 

* Order LeguminosaB. 



6 INTEODUCTION. 

It was not, however, till the spring of 1865 that I was able to carry out this project. 
The careful researches of Wollaston and others had brought to light many interesting 
forms from Madeira, the Canaries, and Cape Verde Islands ; but the Azores had been 
very imperfectly explored, and it was with the idea of giving a more satisfactory 
account of the natural history of these islands, and to trace their relationship to the 
neighbouring Archipelagos, that I decided to investigate their fauna. Oranges from 
St. MicJiael's then formed almost the only trade with England, and in connection with 
the business large numbers of schooners were employed, but with this exception 
there was no regular, direct communication with England. A small steamer, which 
carried the mails, left Lisbon once a month for the two nearest groups of islands, but 
rarely visited the two outer ones. Accompanied by my brother, Captain Temple 
Godman, I took a passage in this vessel, and shortly after our arriviil at St. Michael's 
we were joined by Mr. Brewer, a well known colcopterist whom I had engaged for the 
])urpose of collecting. Interest in island faunas had been much stimulated by the 
])ublication of Darwin's "Geology of the Voyage of the 'Beagle'" and other works on 
the subject. The question was whether the Azores had in former times formed part 
of a continent now submerged, as Professor Edward Forbes believed, or whether they 
had been thrown up from the sea bottom by volcanic agency. After spending four 
months on the islands, during which time 1 visited all except Santa Maria, I came to 
the conclusion that they had always been volcanic islands, and that they derived their 
fauna and flora from neighbouring lands. In 1870 I published a small octavo book 
entitled 'The Azores,' enumerating the plants and animals as far as then known, and 
setting forth my reasons for the conclusion above stated. 

The visit to the Azores was followed in 1873 by an expedition to Madeira and the 
Canaries, in order to compare their respective faunas more criiically, but unfortunately 
the regulations in respect to quarantine were then so stringent that, apart from Madeira, 
my investigations were limited to the island of Teiieriffe. There can, I think, be no 
doubt that the conclasi(>n I had already formed with regard to the Azores was fully 
borne out in these islands also. 

In the meantime, Salvin was residing at Rotherham, Yorkshire, looking after some 
ironworks in wliich he was financially interested; but this was not a congenial employ- 
ment, and he soon gave it up. 

In 1865 he married Caroline, daughter of J. Whitaker Maitland, of Loughton Hall 
in Essex, and they lived for some years at 23 The Boltons, South Kensington, which 
became for a time the headquarters of our Museum. 

Although both Salvin and 1 had jointly collected ever since our iinacrgraduate days, it 
was not until the material was housed at S. Kensington that we really did serious work 
too-ether ; but from thence onwards we spent the greater part of the week in London 
arranging our collections, publishing papers on them, and attending the meetings of 
various scientific societies of which we were both members. 



. INTEODUCTIOX. 7 

Salvin's fourth and last visit * to Guatemala was made in coojpany with his wife, to 
vhose skilful brush we are indebted for the coloured plates of the plants figured in the 
Botany of the ' Biologia.' 

They sailed in a Royal Mail Steamer in April 1873, touching at St. Thomas and 
Jamaica ; then crossing the Isthmus of Panama they reached the City of Guatemala early 
in June. Proceeding thence to Dutfias, which became their headquarters for some 
months, Salvin occupied Limself in collecting in the forests on the mountain slopes. 
Together they ascended the crater of the Volcan de Fuego, and a few days later that 
of Acatenaugo. Leaving Dueiias for Atitlan they made the ascent of the peak from 
Santa Lucia on Jan. ]7th. Subsrquently thry visited Mazatenango, ihe coffee estate 
of Las Nubes (Cerro Zunil), Qu(-zaltenango, the Lake of Atitlan, Solola, Pantaleon, and 
San Geionimo, and after a short stay at the last named place the journey was continued 
to Coban. Having revisited the Capital they left Guatemala in March 1874. Again 
crossing the Isthmus of Panama, they sailed for the United States and visited the 
museums of New York, Boston, Philadelphia, and Washington, and made the acquaiir- 
tance of the leading scientific people, returning to England on June 4th, 1875. 

In the autumn of the same year, Salvin was appointed to the Curatorship of the 
Strickland Collection of Birds at Cambridge, which necessitated his residence at the 
University. On his giving up the house in South Kensington in 1 873, it had become 
necessary to find fresh quarters for our museum and library, which now occupied a 
considerable amount of space, and we took for this purpose a house in Tenterden 
Street, Hanover Square, to which they were transferred. The building being rather 
larger than we requiied, we shared it with some of our ornithological friends, of whom 
Lord L'lford, Dresser, and Seebohm were among the number. After the evening 
scientific meetings of the Zoological Society, the offices of which were then in Hanover 
Square, our rooms became a favourite social resort of ornithologists, and many 
pleasant and instructive hours were spent there. 

During his stay at Cambridge, Salvin came frequently to Tenterden Street and 
worked at the collections with me, and continued so doing until the death of his father 
in 1880, when he succeeded to his property at Fernhurst, Sussex, where he afterwards 
resided, continuing, however, his work in London as before. In the autumn of 1878 
we moved our museum and library to 10 Chandos Street, Cavendish Square, and 
here they remained till after Salvin's death. In 1907 the house at Chandos Street 
was given up and the library transferred to 45 Pont Street, S.W., while the collections 
still remaining in our possession were handed over to the British Museum. 

In addition to the material obtained during our various visits to Central America 
and that sent us by the natives we had trained, we found it necessary, for the suke of 
comparison, to acquire a more thorough knowledge of the South American fauna, and, 

• In the notice of liis journej-s given in tbe Introduction to ti.e ' Aves ' (i). viii) the second expedition, 
1859-1860, was ntt mentioned, and the last visit was incorrt-ctly stated to have heeu made in 1867. 



8 IXTEODIJCTION. 

with that view, emphjyed various expert collectors, whose names are recorded in the 
body of the work, to visit special localities in Colombia, Ecuador, Peru, Guiana, and 
other places in South America. We continued to receive consignments from them for 
several years without having any idea of publishing a connected account of the results, 
and a very large amount of material, especially amongst the birds and insects, was tlius 
accumulated. It was not till the year 1876 that it was suggested that the 'Biologia' 
should be undertaken, and three years later (September 1879) the first part appeared. 

In the meantime, various collectors were sent by us to Guatemala and other parts of 
Central America. 

Our method of publication was to bring out six quarto parts a year ; each part to 
contain twelve sheets made up of various subjects with six coloured plates, the plates and 
letterpress so numbered and paged that the parts might ultimately be broken up and 
bound together in their respective volumes when completed. In this way it was possible 
to keep several subjects in progress at once, and the plan answered well. We were, 
however, unable to adhere to the original scheme of completing the work in GO parts, 
owing to the ever increasing amount of material received from our collectors — an 
amount so great that 215 Parts of Zoology alone have been required, the dates of 
i«sue extending over a period of 36 years. Even now some families of Insects, the 
Crustacea, &c., have not been dealt with— though this is chiefly attributable to the 
fact that no experts on these subjects were available. 

On arrival in England, the various consignments were opened, every specimen 
labelled with its exact locality, and the name of the collector attached. The animals 
sent were then sorted into their respective orders and families, and as occasion offered, 
handed over to specialists to be worked out. It was obviously impossible that we 
could undertake every subject, but the birds and the butterflies we set aside for our 
own share of the work. The names of the various authors who kindly helped us will 
be a sufficient indication of our good fortune in securing the services of so many 
eminent men, all of whom joined the enterprise with great spirit. 

In 1886, in company with Mr. II. J. Elwes, I visited Bombay, Simla, Delhi, Benares, 
Calcutta, and Darjeeling; thence crossing the Rungeet River by a swinging bamboo 
suspension bridge we entered native Sikkim, collecting plants, birds, and butterflies 
en route. At Darjeeling I purchased a large collection of butterflies, made by 
Lidderdale, which at the time was considered very representative of the Northern Indian 
fauna. Starting again from Darjeeling we made a second trip, this time on horse- 
back, travelling by the Government road constructed by Sir Richard Temple along 
the Nepaul ridge till we reached Falute (16,000 ft.), and there passed the night in 
a fairly comfortable bungalow. Next morning we witnessed a magnificent sunrise and 
obtained splendid views of Mounts Everest and Kinchinjunga rising from the mighty 
Himalayan range. I also visited Madras and Southern India and Ceylon before 
returning home. 



INTRODUCTION. 9 

In the autumn of 1887, having been ordered abroad for the benefit of my health, 
I decided to visit Mexico, as in working out the Central American fauna, especially 
the Birds, we found ourselves more deficient in material from that country than from 
further south. Crossing the Atlantic to New York I took the train to Mexico City, 
then a six days' journey, entering the Mexican Republic at El Paso (3700 ft.) on the 
Rio Grande. From this river the land gradually rises and spreads out into the great 
plateau of Central Northern Mexico, bounded on the east and west respectively by 
ridges of high mountains covered with pines, and falling abruptly on the east to the 
Atlantic and on the west to the Pacific. The rainfall being chiefly on the two coasts, 
the plateau is extremely dry and arid, and the vegetation consists largely of cacti, 
yuccas, agaves, and mesquite {Prosopis), with willows and poplars along the margins 
of the few water-courses or lagoons. The early part of the journey was passed during 
the night, but next morning it was evident that a considerably higher elevation had 
been attained, and on reaching Zacatecas (8000 ft.), a large mining district, the aspect 
of the country was very desolate and unpromising for natural history purposes, and 
continued much the same for some distance, but improved a little nearer to the Capital. 
After spending a few days in Mexico City in making necessary preparations, I was 
joined by W. B. Richardson, an American bird-collector, and taking him with me 
started for the Atlantic coast. The first part of the journey, still on the high plateau, 
was through fields of ' agave ' grown for the production of the fermented drink 
called ' pulque,' so much beloved by the Indians. In about six hours we reached 
Esperanza, in the State of Puebla, at the Eastern edge of the plateau ; here the train 
enters the wonderful gorge by which it descends to Orizaba, Cordova, Atoyac, and 
Vera Cruz. The scenery at once changes, the vegetation becoming luxuriant on 
entering the region of the rainfall. 

Before reaching Orizaba we crossed and recrossed the gorge by a series of viaducts 
amongst palms, tree ferns, and tropical plants, loaded with orchids and tillandsias. At 
Orizaba we spent a few days collecting, and I was there joined by Mr. and Mrs. H. H. 
Smith, who went with me as far as Atoyac, where they remained for some time before 
crossing to the Western side of Mexico. The Smiths had previously been in Brazil, 
where they made large collections of insects, which are now in the Pittsburg Museum. 
Mrs. Smith was also skilled in skinning birds which were shot and brought to us by 
the Indians, and through her we made many additions to our store of ornithological 
treasures. Leaving the Smiths at Atoyac, a village at the foot of the steep descent from 
the plateau — still, however, about 1500 feet above the sea and about fifty miles from 
the coast, — Richardson and I continued our journey to Vera Cruz, the land gradually 
sloping down to the Atlantic and forming a savanna or plain of sandy ground, sparsely 
covered with grass and scrub. After spending a few days in collecting, chiefly to 
the north of the town, we took the mule tram to Jalapa, which being on higher ground 
reaps the benefit of the rainfall and the vegetation is far more luxuriant. At this 

BIOL. CENTR.-AMER., Introd. Vol., January 1015. c 



10 HS-^TEODUCTION. 

place I engaged Mateo Trnjillo, a half-breed Indian, who accompanied me during the 
greater part of the time I was in Mexico and proved a very skilful collector. He was 
a first rate climber, and amongst other things made a considerable collection of the 
frogs, newts, and insects which inhabit epiphytical Bromelias growing on the trees in 
the neighbourhood of Jalapa. We made an excursion to Misantla and Papantla, on 
the low ground near the coast at the foot of the mountain range, which proved to be 
very good collecting ground and added considerably to our birds and insects. On 
leaving Misantla we returned to Jalapa and over the Cofre de Perote to Esperanza, and 
thence by train to Mexico City. From the Capital I made two expeditions, the first 
in company with Mr. Flohr to the pretty town of Cuernavaca, and thence to the caves 
of Cacahuimilpa, where he hoped to have found some blind insects. The second and 
also interesting expedition was to Morelia and Lake Patzcuaro, where 1 added to the 
collection of Birds, but it was attended with no very valuable results. 

On returning to the Capital I next visited Yucatan, crossing the Gulf of Campeche 
from Vera Cruz to Progreso by steamer, thence to Merida by train, where the railway 
then ended. My first object was to visit the well known naturalist Dr. Gaumer, who 
kindly accompanied nie to the celebrated ruins of Ticul and Uxmal, a distance from 
Merida of about forty miles, which journey was performed on horseback. The low 
forest through which we passed is said to be still frequented by the beautiful ' Pavo 
real ' [Meleagris ocellata), but it has become so rare in the neighbourhood that we 
ourselves did not meet with it. Leaving Yucatan I went back to Mexico City, where 
I was joined by Mr. and Mrs. Elvves, and together we went to Jalapa, thence riding 
across the country to Cordova, a most delightful journey of three days, and obtaining 
magnificent views of the Volcano of Orizaba, with its snow-clad peak. We continued 
our journey to Puebla and Mexico City, and thence to Amecameca at the foot of the 
volcanoes Iztaccihuatl and Popocatepetl; the latter we ascended as far as the pine belt, 
shooting many interesting birds and collecting plants and insects. We left Mexico in 
the spring of 1888, having been absent from home about five months, and returned to 
England vid California and New York. 

For several years after my return from Mexico, Salvin and I continued diligently 
to work out the material on hand and the ever increasing amount sent over by our 
collectors. Salvin's failing health finally obliged him to relax his eff'orts, and though 
he still came to London as formerly, he was unable to take the same active part 
in the work and the diflaculty of concentrating his attention on any one subject 
became increasingly great. He died suddenly at Hawksfold, Fernhurst, Sussex, 
June 1st, 1898, leaving me alone to complete the ' Biologia.' 

The severance of a friendship such as ours had been for forty-four years was a terrible 
blow to me, for we were more intimately connected than most brothers, and, besides 
the personal loss, I missed his knowledge and experience in all things connected with 
our book. At the time of Salvin's death, 141 Parts of Zoology (completing 13 volumes), 



INTKODUCTION. 11 

the whole of the Botany (5 volumes), and nine Parts of Aicha?ology had been issued: 
since then 74 Parts (142-215) of Zoology, completing 39 more volumes, and 8 Parts 
of Archaeology have been required, to bring these subjects to a conclusion. It 
was with a heavy heart that I took up my pen again. With the assistance of 
Mr. G. C. Champion I continued Vol. II. of the Rhopalocera and finished it in 1901, 
and in 1904 Vol. III. of the ' Aves' with the help of Dr. R. B. Sharpe, of the British 
Museum, was also completed. 

Salvin had made a special study of the Procellariidse, a very difficult group of birds, 
and for a long time we had missed no opportunity of adding specimens of Petrels to 
our collection. After writing the catalogue of this family for the British Museum, he 
intended to publish an illustrated monograph on the subject, and for that purpose 
40 plates had already been drawn by Keulemans. It remained for me to carry out 
his intention, and having again sought the aid of Dr. R. B. Sharpe we completed 
the work in 1910, enumerating 123 species, and illustrating them with 106 coloured 
plates. 

It will be seen that the travels of Salvin and myself covered only a comparatively 
small part of Mexico and Central America, but we employed a considerable number 
of expert collectors to travel in districts we had not visited, and they continued to send 
us the results of their labours for some years after we had left. Amongst them 
must specially be mentioned W. H. Richardson, Mr. H. H. Smith, and Mateo Trujillo, 
all of whom accompanied me while I was in Mexico, and Lloyd and Armstrong, who 
devoted their attention particularly to the Northern provinces of that country. 

In working out the Mexican Birds we found ourselves hampered for want of 
an authentically named collection of North American species for comparison, which 
did not exist in Europe at that time. In order to remedy this, I acquired the Henshaw 
collection, numbering 13,326 specimens, and this was rendered still more valuable 
through the courtesy of the authorities at the United States National Museum, 
who allowed Mr. Ridgway, the highest authority in America, to go through and 
verify all the names on tlie labels attached. I likewise purchased 2500 carefully 
named birds from Florida from Mr. W. E. D. Scott, 321 named specimens from 
California, Texas, &c., from Mr. C. K. Worthen, and a series of Mexican birds from 
Senor F. Ferrari-Perez ; and these collections proved of great advantage to us. 

In a similar manner we dealt with the insects, &c. I bought H. W. Bates's 
collection of butterflies, including those he obtained from the Amazons, as well as 
that of Herbert Druce, containing the ' Kaden ' types ; the first set of the extensive 
series of Mexican and Central American Coleoptera amassed by A. Salle (including the 
types of several of the older authors, and some thousands of specimens found by 
himself or by M. Boucard). I also acquired the general collection of Heteromera of 
F. Bates (22,390 specimens); a portion of Dr. J. S. Baly's collection of Phytophaga ; 
the Janson coUectiim of Elaterida; (25,000 specimens); various Reptilia, Coleoptera, &c., 

c2 



12 INTEODUCTION. 

collected by A. Forrer in N.W. Mexico and the Tres Marias Islands ; a second set of 
the very large number of Coleoptera obtained by C. T. Hoge in his two expeditions 
to Mexico, the first set now in the Berlin Museum, having been retained by his 
employer, Mr. Flohr ; collections of insects from Messrs. Becker, Biolley, Blancaneaux, 
Conradt, Gaumer, Janson, Lankester, Morrison, Staudinger, Underwood, Van Patten, 
Wittkugel, &c. In addition to this material, we had, of course, the whole of that 
procured by our other collectors, E. Arce, G. C. Champion, and H. Rogers. Further 
details are appended on pp. 44, 45. 

All the insects from Mexico and Central America, the Salle and Janson collections 
of beetles, our own general collections of birds and butterflies, and the Henshaw 
collection of birds, have been presented by us to the British Museum, and are being 
gradually incorporated with the National Collection. 

The various accessions are enumerated in detail in Vol. II. of the ' History of the 
Collections contained in the Natural History Departments of the British Museum' 
(1906) and in the subsequent Annual Reports of that Institution. The first instalment 
of Neotropical birds (50,120 specimens) was presented in January 1885, and other 
instalments followed from time to time till the whole of them became the property 
of the Nation. Amongst the insects, up to 1906, the total number of specimens 
given in the ' History ' is as follows : Coleoptera (85,920), Lepidoptera Rhopalocera 
(17,829), Lepidoptera Heterocera (12,883), Diptera (17,525), Hymenoptera (10,004), 
Rhynchota Heteroptera (5543), &c. These figures do not include the Rhynchophora 
or weevils (22,793), the Staph ylinidse and water-beetles (9474), the Odonata (3000), 
the Rhynchota Homoptera (5509), the supplementary unworked parasitic Hymenoptera 
(6293), &c. From 1906 onwards the remaining collections have been handed over 
to the Museum as soon as the enumeration of the species was completed ; that of 
the Coleoptera was finished in 1911. Our own general collection of butterflies 
probably included nearly 100,000 specimens, and the beetles alone from Mexico and 
Central America perhaps double that number. Besides these a considerable number 
of mammals, reptiles, fish, &c., of which no account was kept, were presented to the 
National Museum. 

F. D. G. 



[ 13 ] 



PHYSICAL FEATURES ETC. OF THE AREA TREATED. 



TuK area treated in the * Biologia ' includes the whole of Mexico south of the 
Rio Grande as far as El Paso, thence to the Gila River, and following it as far as 
the Gulf of California (but excluding Baja or Lower California). The distant 
Revillagigedo Islands have been added, owing to certain species of sea birds being 
common to these islands and theTres Marias on the western coast of Mexico. Further 
south we include British Honduras, Guatemala, Salvador, Honduras, Nicaragua, Costa 
Rica, and Panama, which collectively are spoken of as Central America. The country 
stretches in a south-easterly direction, having a width in the north of about 1140 statute 
miles and gradually decreasing till at the Isthmus of Panama it does not exceed 
45 miles, and the land only attains an altitude of 300 feet (tierra caliente). In 
shape Mexico and Central America have been likened to a cornucopia, whicli 
collectively they somewhat resemble. Of the pliysical aspect of each country I 
propose giving an outline, but as the travels of Salvin and myself covered only a small 
portion of the area, I shall supplement the account with extracts from other writers; 
moreover, upwards of fifty years have elapsed since I was in Guatemala, and doubtless 
great changes have taken place in the interval throughout the whole of Mexico and 
Central America, partly owing to the extensive destruction of forest for the purpose 
of cultivation, the construction of railways, and the wanton devastation of large tracts 
by fire. Thus various places which in our time were good collecting grounds are no 
longer so, and many species of both animals and plants must either have migrated 
or become extinct. Earthquakes, too, have done much to alter the configuration of 
the land, as well as the nature of the vegetation. 

The country is divided by the natives into three zones — the * tierra caliente,' ' tierra 
templada,' and 'tierra fria ' (or hot, temperate, and cold climates respectively). The 
tierra templada corresponds on an average with an elevation of 3000 to 5000 feet, but 
the natives of the Mexican State of Vera Ciuz draw this imaginary line at a level very 
different from that used by the people on the western slope of Mexico. For instance, 
Chilpancingo at 4000 feet in Guerrero has no tropical vegetation and the climate is 
much cooler than in the State of Orizaba at an almost similar altitude on the eastern 
slope, which is in every sense subtropical. 

The difi'erence is due to the greater rainfall on the Atlantic coast, for the prevailing 
wind in ])assing over the ocean is charged with humid air, and when driven by the 
mountain ranges into a higher and colder elevation, the moisture is then precipitated 
as rain, thus inducing a much more vigorous vegetation than is found on the Pacific 
slope. 



14 PHYSICAL TEATUEES ETC. 

MEXICO. 

The general aspect of the Mexican plateau, as far south as the City of Mexico, has 
been previously alluded to in the account of my journey there in 1887-1888 {antea, 
pp. 9, 10). On each coast, between the mountain ranges and the sea, there is a com- 
paratively narrow strip of low land, producing a tropical vegetation, which on the 
Atlantic extends a little to the nortli of Tampico, where the forests with epiphytical 
orchids suddenly disappear; while on the Pacific the low land terminates a little above 
Mazatlan, where the coco-palm ceases to grow, but the vegetation generally is less 
luxuriant on this coast than on the Atlantic, The plateau itself is arid in consequence 
of the rainfall being precipitated on the two coastal ranges, and produces but a scanty 
vegetation, consisting chiefly of Yuccas, Agaves and Cactaceae, and Mesquite [Prosopis). 
The margins of the few existing streams or pools produce a few scattered willows and 
poplars, and the whole country presents a desolate appearance. The plateau is indeed 
a continuation of that of Arizona and New Mexico, and though it is depressed in the 
valley of the Rio Grande at El Paso, where the railway crosses, it is still at an altitude 
of 3700 feet above the sea. Southward it mounts considerably higher, and on 
reaching Zacatecas, the highest point on the railway leading to the Capital, there 
is an elevation of 8000 feet. The City of Mexico is situated in a valley surrounded by 
ranges of hills clothed towards their summits with pine trees, and reaching an elevation 
of about 10,000 feet, while to the south-east are the two lofty volcanoes of Popo- 
catepetl and Ixtaccihuatl, the tops of which reach above the snow limit. Numerous 
streams descend from the mountains and empty themselves into an alluvial valley wich 
svy'ampy meadows and form several large lakes, of which Texcoco, Chalco, and 
Xochimiico are the most important. It was on Texcoco that the old Aztec Capital 
was situated ; the houses were built on piles, and the city, which was approached by 
a causeway, was destroyed by Cortes at the time of the Spanish conquest in 1519- 
1521. The modern town was placed by the conquerors on the swampy ground about 
two miles from the eastern margin of the lake, but notwithstanding the altitude it 
became one of the most unhealthy capitals in the world, for owing to the frequent 
rising of the level of the lake, the town was constantly inundated. Eventually a great 
drainage scheme was inaugurated, and for 150 years vast numbers of natives were 
employed in cutting a huge dyke for the purpose of draining the overflow of the lake ; 
but the sodden ground had become so impregnated with sewage that the mortality 
was still extremely high, and it was not till 1900 that President Diaz finally completed 
the extensive drainage system which now renders the city a comparatively healthy 
resort. There can be but little doubt that in former times one large piece of water 
covered the whole area of the three lakes, and a map published about 1628 shows 
Chalco united with Texcoco, when the surface of the two lakes was far greater 
than it is at present. Owing to the drainage system so recently completed, there are 



MEXICO. 15 

Tiow several separate lakes draining from one into the other, and finally ending in 
Lake Texcoco, which is greatly reduced in size ; the water of this lake is brackish, 
while that of the other is fresh. 

An account of my journey from Mexico City to Orizaba has been already given 
(p. 9), so, after stating that I found myself on a limestone formation surrounded by 
rich vegetation, with the high peak of Citlaltepetl towering above, I will refer to 
Dr. Gadow for his description of the ascent of this volcano in company with his wife 
('Through Southern Mexico,' chap. iii. 1908). He gives in some detail the change 
of climate at various altitudes, and its consequent influence on the Flora and Fauna, 
whicli is of special interest here, as conditions somewhat similar prevail on nearly all 
high mountains in the tropics. 

Starting from Orizaba, Dr. and Mrs. Gadow camped near the village of Xometla at 
an altitude of 8600 feet, where they remained for a few days exploring the neighbour- 
hood. On their way there they crossed a deep limestone gorge, with fertile vegetation 
consisting of a species of Platanus, magnolias, crotons, and various kinds of oaks, 
most of these supporting a luxuriant growth of bromelias, ferns, selaginellas, and 
orchids, interspersed with lichens and tillandsias which proved to be " hotbeds 
of life." Northward the open slopes were covered with pasture and clusters of trees 
and shiubs, including mimosas, acacias, yuccas, plane-trees, and bamboos. Here was 
reached the upper limit of coffee and cotton plantations, while in the damper ravines 
tree-ferns were plentiiul. Higher up, at the level of the central plateau, maize fields 
became scarce, and tree-ferns and datura disappeared. At this altitude a great 
change in the vegetation takes place ; there is now but little trace of tropical plants, 
and the climate is temperate, moist and fertile, coinciding with the cloud belt. The 
vegetation near the camp consisted chiefly of pines. P. montezumce and P. liophyUa, 
with open spaces bordered with deciduous and evergreen oaks, arbutus of two species, 
alder and Fuchsia microphylla, with tillandsias in abundance. The larger tillandsias 
occur up to a level of 9600 feet, where they suddenly disappear, and mistletoe then 
takes their place on the trees. In the ' barrancas ' or gorges are high trees covered 
with creeping aroids and lianas hanging from the branches with abundance ot 
bamboos and maiden hair fern. Here bird life was almost absent, and only a few tits. 
a tree-creeper, a woodpecker, and some blue jays were recorded. A few small 
mammals were not uncommon, and the armadillo, which is fairly plentiful in the 
lower and tropical country, still exists at an elevation of 8000 feet. Several species of 
amphibia and reptiles are also found at this altitude, living chiefly in the bromelias 
and other epiphytical vegetation. They have either no lungs or only tiny vestiges 
of them, respiration being chiefly carried on through the moist skin. One genus, 
Spelerpes, has a wide distribution in Mexico, and S. orizabensis, which leads only 
a partially arboreal life, ascends to an altitude of 12,500 feet, llylodes rhodojjis, 
which leads the life of a tree-frog, occurs at 10,000 feet, while it also inhabits the 



IG PHYSICAL FEATURES ETC. 

low hot country of Vera Cruz. Snakes of the genus Crotalus are common in tlie 
neighbourhood of Orizaba, but disappear entirely in the wet and cloudy zone about 
Xonietla, being again represented by a small species with a poor rattle at an elevation 
of 13,000 feet. Of harmless snakes, only Tropidonotus scalaris was met with, a 
representative of a typically northern genus. A small lizard, Sceloporus microlepidotus, 
also has a great vertical range from the hot plains of Oaxaca to the upper tree limit of 
Citlaltepetl. 

At their highest camp, 12,500 feet, were stunted pines and juniper, while at their 
feet grew asphodel and tussock grass, which continued to an altitude of nearly 
14,000 feet. At 14,400 feet they reached what appeared to be permanent patches of 
snow, but the white summit of the peak, which from this side they found impracticable 
of ascent, was still 4000 feet above them. 

At a later date Dr. Gadow, however, ascended the peak itself, an elevation of 
18,200 feet, approaching it on the north-east side from La Barbara. 

Proceeding in a south-easterly direction from Cordova (2700 feet) there was dense 
tropical vegetation on limestone formation all along the foot of the slopes. Eastward 
the country showed open savannas, followed by lagoons, swamps, and low forests 
in the neighbourhood of Agua Fria. The Rio Papaloapan with its many tributaries 
carries an enormous volume of thick yellow water, and for months the low ground 
is to a great extent submerged. Further on. Dr. Gadow describes low undulating 
grazing land, until he reached the railway which crosses the Isthmus of Tehuantepec. 

The eastern slope is clad with humid evergreen forests, but farther west the line 
crosses a ridge at an altitude of rather less than 1000 feet, and passes through dense 
tropical forests with occasional open patches, but on the Pacific coast a drier type of 
vegetation prevails. From Salina Cruz and Tehuantepec north-westwards to the City 
of Oaxaca the land again ascends towards the high plateau. The hills at first are 
covered with low vegetation, until at 3000 feet pines and oaks appear, but nowhere is 
there continuous forest. The general character of the country is dry. Deep gorges 
and sandy river beds alternate with an intricate system of hills and patches of 
xerophile vegetation. Looking south from an altitude of 5300 feet, the tierra calientc 
appears to be densely covered with wood, while to the north is seen a fiat, almost 
treeless plateau, with here and there outcropping barren ridges of volcanic nature, or 
with wooded slopes of Palaeozoic formation. 

The whole of the Western or Pacific slope is much drier than that of the Atlantic, 
and the vegetation is consequently less dense. Pine trees descend to about 2000 feet, 
and cacti prevail. 

It was on this occasion that, in company with Mr. Julius Flohr, I made a three days' 
expedition on mule-back from Mexico City to the Cave of Cacaliuimilpa, stopping 
at the pretty little town of Cuernavaca en route, from which a fine view of the 
western slope is obtained. 



MEXICO. 17 

The caves, though of considerable size, were on the whole rather disappointing, 
and the only animals seen were innumerable bats, which tainted the whole atmosphere ; 
as regards other living creatures, even after a diligent search, neither Mr. Flohr nor 
myself succeeded in discovering any traces of insects. A Lepisma and a beetle are, 
however, reported as having been found by other. collectors. 

Gadow's account of the Rio Balsas or Mescala basin informs us that it is bordered 
on the south side, parallel with the Pacific coast, by a long high range of mountains, 
attaining an altitude of 10,000 feet, densely wooded and intersected by deep gorges, 
while the river beds, which form the only available roads, occasionally widen into 
meadows. 

At Chilpancingo there is a wind swept, shallow depression of cretaceous formation, 
surrounded by sparsely wooded hills with meadows on the top of the ridge. To the 
west the slopes of the Sierra Madre del Sur are covered with rich forest growth : oaks, 
dwarf palms, and pines abound, higher up oaks, pines, and arbutus, and, finally, pines 
alone, form dense high forests ; while in the gorges, especially within the cloud-belt, 
most luxuriant undergrowth prevails. Omilteme (7100 feet) where many specimens 
were obtained, is situated in these mountains. 

At Cumbre de Los Cajones, a pass of 3500 feet marks the beginning of volcanic 
formation. The typical ' tierra caliente,' with an essentially tropical flora and fauna, is 
found on the southern slope of the main ridge, coinciding with what is officially known 
as La Costa. The upper limit may be put at 1000 feet, but the country loses its 
tropical character on the ridges, which rise higher than 1500 feet. 

From Coquillo to Chacalapan (700 feet) there is tropical life, and from thence to the 
coast across lower ridges, the subsoil consists of gneiss and granite in rapid decom- 
position, while wooded ground and open pastures are also found. The rivers during 
the winter season are frequently dry, but near the granite bound coast are lagoons 
mostly containing pure water ; a broad belt of almost impenetrably high forest extends 
in places to the sea. Mangrove swamps are permanent features of the landscape, but 
in the rainy season many parts of the forests are inundated, and over nearly the whole 
of the coastal district a dense mass of tall herbs usurps the place of brushwood in the 
forest belts. 

Throughout the State of Guerrero large collections, chiefly consisting of birds and 
insects, were made for us by Mr. and Mrs. H. H. Smith, who spent a considerable 
time in the district, paying special attention to the fauna during the time 1 employed 
them in Mexico. 

We have no detailed account of the States of Uurango, Sinaloa, and Sonora, but the 
general aspect is barren and rocky, and although I sent one or two collectors to 
those districts, I gained no accurate information as to the physical features. 



BIOL. CENTR.-.\MER., Introd. ^'ol., January 1915. 



18 PHYSICAL FEATURES ETC. 

EEVILLAGIGEDO ISLANDS. 

Neither Salvin nor I were able to visit these islands, and the only information 
respecting the physical features and distribution of the avifauna is from the 
account given by Mr. A. W. Anthony in the 'Auk,' xv. pp. 311-318, after his visit 
in 1897. 

The Revillagigedo Islands lie to the S.W. of Cape San Lucas, Lower California, and 
consist of a group of four islands, at some considerable distance apart, but all are of 
volcanic origin and in general appearance extremely rough and broken. Socorro, the 
largest of the group, is about 240 miles south-west of Cape San Lucas and about 285 
miles to the westward of Maria Madre, the largest island in tlie Tres Marias group, 
off San Bias. It is estimated to be about one hundred square miles in extent, and rises 
to a height of 4000 ft. in the centre, where there is an extinct volcano. The greater 
part of the island is covered with a dense mass of undergrowth which it is almost 
impossible to penetrate, especially on the north and north-west, or weather side. 
Trees are abundant there, but do not exceed forty or fifty feet in height, though 
usually covering a considerable area with their spreading branches. On the south 
and east the trees are mostly confined to the canons, where they are smaller than on 
the northern slope. 

Clarion Island lies approximately about 200 miles westward of Socorro and some- 
what further south, and has little in common with the other islands, either in fauna 
or flora. Its length is about five miles, and width one mile, the ground rising about 
1500 feet above the sea. A few low trees or shrubs, the largest not over ten feet in 
height, are scattered along the main plateau, and in a few places extend to the level 
ground on the south side, which lies between the Mesa and the coast. Nearly the 
whole of this flat ground is covered with a dense growth of cactus {Platopuntia), 
over which has grown a mass of vines, and a passage through this belt can only be 
made by the diligent use of a bush knife. At a short distance from the beach were 
found two small shallow ponds, which contain water during the rainy season only, 
but as the high tides evidently overflow the barriers and flood them with sea 
water, it is doubtful if they are ever otherwise than brackish. At the time of 
Mr. Anthony's visit, the ponds had been dry for some months, and no water was 
found upon the island. 

As might be expected from the position and vegetation of Clarion, the birds were 
quite different from those on Socorro and San Benedicte. The only land bird common 
to any two islands was the Raven, but on Socorro it was not seen. 

San Benedicte is a small island about three miles in length with an average width 
of half a mile, and lies 35 miles north of Socorro. There is little vegetation, but 
wherever sufficient soil is found, there is a heavy crop of coarse grass growing five or 
six feet in height, and rendering progress extremely difficult. The barbed seeds 



EEVILLAGIGEDO AND TEES MAHIAS ISLANDS. 19 

penetrate a man's clothing, and were found to be even more unpleasant than the 
cactus thickets on Clarion Island. 

Roca Partida, or Divided Rock, lies 65 miles north-west of Clarion, and is the 
fourth of the group. It is of considerable elevation and has the appearance of a ship 
under sail. 

The number of species of birds observed by Mr. Anthony are as follows : — Socorro 
(24 — 13 of which were generally distributed land birds), Clarion (17 — 10 of which 
w^ere land birds), San Benedicte (11 — including 3 land birds). 



TRES MARIAS ISLANDS. 

As in the case of t!ie Revillagigedo Group, neither Salvin nor I visited the Tres 
Marias Islands ; but Alphonse Forrer collected in Maria Madre on our behalf and 
obtained a large number of specimens. For a description of the physical aspect of 
the islands we are indebted to Mr. E. W. Nelson, who wrote a ' General Account 
of the Tres Marias Islands with Reports on Mammals and Birds,' supplemented 
by Notes by other authors on the Reptiles, Crustacea, and Plants (North American 
Fauna, No. 14, 1899). 

The islands are situated about 6-5 miles off the west coast of Mexico from San 
Bias, between lat. 21° and 22° and long. 106''-107°, and consist of a group comprising 
Maria Madre, Maria Magdalena, Maria Cleofa, and San Juanito, arranged in a north- 
westerly and south-easterly direction. 

About 20 miles from the mainland lies the small island of Isabel, about 1 mile 
long and 150 feet high. It chiefly consists of the remains of an old volcano, and 
a small crater still occupies the centre. There is evidence of its having once been a 
much larger island, though now apparently sinking. The soundings in the channel 
between the islands and the mainland gradually deepen to nearly 300 fathoms, but 
west of the group the sea bottom falls rapidly to over 1500 fathoms. 

Maria Madre, the largest island, measures 8 by 15 miles, and rises over 2000 feet 
above the sea. The interior is occupied by a mountain ridge extending almost the 
whole of its length, but descending to a gently sloping area at each end. The eastern 
side has the longer slope, while the western or seaward face is much more abrupt, 
thus corresponding with the formation of the mountains paiallel to the coast on the 
adjacent mainland. Both slopes are scored at intervals with canons, which usually 
descend in a nearly direct line to the sea, and along the lower slopes of which, Spanish 
cedars and wild figs are grown, certain trees attaining a great size. Generally speaking, 
the forest is low and scrubby near the shore, but increases in luxuriance farther up the 
slopes. In its primeval condition before the advent of wood cutters, a fine example 
of typical forest growth must have been presented here. 

North of Maria Madre, and separated by a channel 4 miles wide and 5 or 6 fathoms 

d2 



20 PHYSICAL FEATURES ETC. 

deep, is an islet 3 or 4 miles in diameter and only about 100 feet In'gh ; there is a 
narrow border of buffs along the northern shore, thus forming an exception to the 
other islands, which are mountuiuous and rise in successive slopes to the culminating 
point near the centre. Here the vegetation largely consists of hush and scrubby 
trees 8 to 15 feet high, with many agaves on the sandy southern end. 

South-east of Maiia Madre, and separated by a shallow channel 8 miles wide, is 
Maria Magdalena, roughly triangular in outline, and 7 or 8 miles across, rising in the 
centre to an altitude of about 1500 feet; south-east again lies Maria Cleofa, the last 
of the group. In shape it is irregularly rounded, and about 3 miles in diameter ; 
the altitude is apparently much less than 1320 feet, as recorded on the charts. 
The channel between the two last named islands is about 12 miles wide and much 
deeper than the others. Maria Magdalena and Maria Cleofa have a central 
mountainous elevation from which canons descend in all directions to the sea. 

The north-eastern points of both islands are low, flat, sandy areas of limited extent, 
•while the western faces are rocky and precipitous. Permanent fresh water is very 
scarce on all the islands. 

"When visited near the end of the long dry season in May 1897, most of the 
herbaceous plants were withered. The general appearance of the vegetation was, 
however, the same as in similar situations on the mainland. The most noticeable 
plants were Spanish cedar (Cedrela), 3 species of wild fig, 2 of Pithecolohium, 5 of 
Solanum, 2 of Ipomoea, a Passiflora, Cassias, Euphorbias, a large Agave, a large Cerens, 
and 2 Opuntias. 

The following is a summary of the species of animals and plants known from the 
Tres Marias in 1897, as quoted by Mr. Nelson : — Land mammals, 11 (7 peculiar) ; 
birds, 83 (24 peculiar) ; reptiles, 18 (1 peculiar) ; freshwater fish, 2 ; freshwater 
shrimp, 1; land molluscs, 6; plants, 136 (12 peculiar). Two species of bats found 
by Forrer were not met with by Mr. Nelson, and he was of opinion that both were 
stragglers from the mainland. 

The relative situation of this group of islands, all with narrow, shallow channels 
between them, shows conclusively that at one time they formed a single island at least 
45 or 50 miles long, and at a still earlier stage they must have been connected with 
the mainland. One of the strongest proofs of this former connection is shown by the 
correspondence between the fauna and flora. The breaking down of the original 
island into several smaller ones and the continuous encroachment of the sea appear 
to indicate that the subsidence is still in progress. The mainland in Tepic near the 
coast was within a comparatively recent period the scene of great volcanic activity, 
and the Tres Marias Islands bear evidence of having undergone various oscillations, 
while the marine deposits of Maria Madre are further indications of the recent 
change. 



YUCATAN. 21 



YUCATAN. 



During my stay in Mexico in 1887-1888, I determined to visit the province of 
Yucatan with the object of exploring the wonderful ruins of Chichen Itza. Taking 
my passage in a steamer from Vera Cruz, I landed at Progreso, a port of Yucatan 
situated on a spit of sand separated from the mainland by marshy swamps, which, 
during the stormy northers that prevail in winter, is occasionally inundated. 

After spending a day or two at Merida, the capital, I visited Dr. Gaumer at Izamal 
about fourteen leagues distant. This American gentleman, long resident in the 
country, had made considerable collections of birds and insects, some of which he 
had previously forwarded to us in England. 

My original intention had been to ask him to accompany me to Chiclten Itza, but, 
owing to the disturbed state of the Indians in the vicinity, he advised me not to attempt 
the journey, volunteering instead to go with me to Ticul and Uxmal, a journey we 
performed on horseback. On leaving Merida we passed through a forest with patches 
of open ground, some of which were cultivated with Indian corn and an agave, 
from which a fibre called ' sisal ' is obtained and exported in large quantities. The 
name ' sisal ' is derived from an old port on the north coast, six leagues from Progreso, 
from whence the fibre was originally shipped. As we proceeded further south, the 
forest trees became larger, but still not of the great size usually found in the tropics. 
The ruins of both Uxmal and Ticul have been very much despoiled, a vast number of 
the stones having been carried away for building purposes, wliile many of the carved 
pieces formed part of the ' hacienda ' at which we resided. From a detailed account 
of these ruins when described by Mr. A. P. Maudslay in the ' Archaeology ' of the 
' Biologia,' it is evident that they were enormously reduced in size since the visit in 
1839 of J. L. Stephens, who published in 1843 an account of them in his 'Incidents of 
Travel in Yucatan,' with admirable illustrations by Catherwood. 

The peninsula of Yucatan is flat and of a recent limestone formation ; there is a low 
range of hills which stretches from a point a few miles south of Merida to the 
neighbourhood of Peto some distance south of Ticul, but nowhere exceeding 500 feet 
in height. 

The coast is very low and swampy, while further inland are forests, which in a 
tew cases have been cleared, but the whole country is very sparsely inhabited. The 
southern part is, so far as I could learn, but little known, but it is said to be largely 
covered with forest and the trees are much finer than those in the north. 

The following description of the country is mainly taken from Dr. Gaumer's notes 
published in Boucard's account of a ' Collection of Birds from Yucatan ' (P. Z. S. 1883, 
pp. 434-439), supplemented by my own observations in 1887-1888. At Tizimin the 
country, like the rest of Northern Yucatan, is on a low level, but to the north east and 
south lie vast forests, for the most part uninhabited since the migration of the Indians 



22 PHYSICAL FEATUEES ETC, 

half a centuiy ago. This region is filled with ruins both ancient and modern, but of 
the former very few of any size remain. Some of the ranchos have, however, been 
re-peopled and most of the birds have been obtained in the vicinity of the clearings. 
At Yok Jonat Ku there is a large forest where the trees are high and the ground 
comparatively open ; here the magnificent turkey Meleagris ocellata is still to be found. 
At one time this bird was distributed all over the peninsula, but owing to the depre- 
dations of the Indians, who esteem it highly as an article of food, it is now almost 
exterminated. 

Lagartos is a sea-port town at the mouth of the river, or more properly an arm of 
the sea bearing the same name, and innumerable streams or — as Dr. Gaumer believed — 
subterranean rivers find an outlet there. The waters are very salt, and in the dry 
season are even more saline than the sea itself. This so-called river is broad and 
shallow, bordered by a dense growth of low bush, behind which lie marshes of salt or 
brackish water, and here in June and July thousands of flamingoes in their finest 
plumage were seen by Dr. Gaumer, while swarms of other sea-birds were always in sight. 
The innumerable hosts of mosquitoes which come with the first rain impeded the 
work here, and the intermittent and ])ernicious fevers render collecting both difficult 
and dangerous. The country generally has no surface water, and the only supply is 
from the Aguadas and Cenotes (Senotes or Jonats), as they are called by the natives. 
Fortunately the Aguadas, which are said to be of natural formation, but which appear 
to have been reconstructed by the ancient Indian races, are very numerous ; they 
consist of a deep excavation in the earth, sometimes circular in form, but giving the 
idea of having been at one time quadrilateral, and from fifty to one hundred feet in 
diameter. They contain water all the year round, though never of any great depth. 
The sides being inclined, they form natural drinking places and are much frequented 
by animals and birds — so much so, that the collector usually obtains a good number of 
specimens in the vicinity. 

The Cenotes are probably natural openings in the earth with steep walls of limestone 
frequently sixty feet high ; they vary in size and shape, but always contain clear, fresh 
water. They are believed to be openings to underground rivers, and are frequently 
found in immense caves with a narrow circular mouth ; at the water's edge there is no 
resting-place and no approach except by the steep sides. Vultures, owls, and similar 
birds nest in the walls. The caves are also frequented by swallows, bats, and motraots, 
and reptiles are said to occur in immense numbers. The water contains numerous 
fish belonging to the Siluridae, and in the shallow open water-holes near the coast 
there is said to be another species belonging to the same group, but Dr. Gaumer 
was, unfortunately, unable to capture a specimen. The distribution of these Siluridaj 
confirms the belief that underground rivers in Yucatan do exist. 

A very interesting description of the climate (in 1878-1879) is given by Dr. Gaumer *, 

• See Boucard, P. Z.S. 1883, pp. 434-402. 



TUCATAX. 23 

who first reached the country in tlie middle of October 1878. The summer rains had 
ceased about ten days previously and the weather had been good, but throughout 
October, November, and December ' norther ' followed ' norther ' every ten to 
fourteen days, accompanied by light drizzling rain wliich lasted generally from two 
to four days with increasing cold. In January there were four moderately heavy 
rainfalls with strong ' northers ' and cold nights. In February there were five 
' northers ' and one with very heavy rainfall accompanied by hailstones of such size 
that they were quickly gathered up and by many people placed in bottles thinking that 
they might so be preserved. From February 26th to May 23rd no rain fell and the 
sky was cloudless for weeks at a time. The heat during the day gradually increased, 
until in April and May it was almost intolerable, and on account of the dry air and 
clear sky tlie radiation was so great that the nights were disagreeably cool, though 
generally so balmy and pleasant. The birds disappeared as the dry season advanced, 
when only a few common resident species round the ranchos and Aguadas were to be 
found. On May 23rd the first summer rains commenced and were followed by daily 
showers at mid-day. All nature changed as if by magic, new leaves grew, and the 
forests were again populated with songsters. In June the rains began at 11 a.m. and 
ceased at 2 p.m. with an almost daily regularity. In July they began at 10 A.At. 
and ceased at 3 or 4 P.M., but never earlier. In August it rained from 10 a.m. till 
nightfall and sometimes later. During these last three months there were from five 
to eight days in each month upon which no rain fell. The heat was almost insup- 
portable, even to the natives, and yellow fever raged in the towns of the interior. In 
September the rains lasted from 8 or 9 a.m. till midnight and not infrequently all 
night, but the weather became milder. Reptiles were almost the only things to be 
found. In October 1879, during the first twenty-seven days, rain fell in torrents and 
almost incessantly, the sun was seen but on four or five days and the stars appeared in 
patches on five nights only, and not five consecutive hours were fine during those 
twenty-seven days. Yellow fever gave place to bilious fever. Insects were rarely seen, 
birds almost entirely disappeared, and any skins were worthless, as either owing to the 
heavy rain or to some other cause the feathers had not yet commenced to grow. 

An account of Yucatan would scarcely be complete without some particulars of the 
distribution of species and a comparison with that of the islands off the coast and in 
the Bay of Honduras, which were visited on our behalf by Dr. Gaumer. In a summary 
of the island-birds examined in detail by Salvin ('Ibis,' 1890, pp. 84-95) he arrives at 
the conclusion that they split up naturally into three groups. Leaving out Meco, the 
exact position of which is uncertain, he considers that those from Holbox and Mugeres 
may be classed together, Cozumel by itself, and Kuatan and Bonaca (Guanaja) by 
themselves, though all show a strong affinity to the birds of the mainland. The two 
last named islands are stated to be very different in their physical features to the 
others mentioned ; they are of high altitude and attain an elevation of 1200 feet, and 



24 PHYSICAL FEATUEES ETC. 

have the upper parts covered with pines, in contrast to the low ground and recent 
coral-limestone formation of the remainder. 

The total number of species obtained was 214, of which 79 are migrants from North 
America, and of the remaining 135 species, 27 are birds which frequent the sea coast. 
The table of distribution of the 108 non-migrating species shows that the affinities of 
the birds of these islands as a whole are largely on the side of those on the mainland, 
the West Indian element being very slight. Northern Yucatan and the islands 
adjoining are separated from Cuba by a depth of over 1000 fathoms, and the Bay 
Islands from Jamaica by over 500 fathoms. Had there been any recent land con- 
nection, a supposition which the similarity of the birds alone would justify, the main- 
land as well as the West Indies Avould hardly fail to show such a connection in a much 
more pronounced manner, and we should not find the strong contrast which exists 
between the faunas of Cuba, Jamaica, and the mainland, but a larger number of features 
in common ; this contrast is still more marked in the Lepidoptera Rhopalocera. 

The alternative supposition to account for the West Indian element in these islands 
is that the birds have reached them at no distant date by flight. AVhen we consider 
that the trade wind blows in the direction of Yucatan and this coast for several months 
in the year, it is scarcely a matter of wonder that some W^est Indian birds stray so far 
west. Cozumel appears to have been separated from the mainland for a considerable 
period, during which time it has received casual immigrants from the West Indies, 
from North America, and from the mainland, some at a distance of time sufficient to 
allow of their modification. There are 159 species of birds, 65 of which are migrants 
and 27 of very wide range. 

My stay in Yucatan was very short, and owing to the state of my health, which 
rendered me unfit for much exertion in a country so little explored, I was able to do 
very little collecting. The specimens acquired, however, Avere, as before, mostly due 
to the enterprise of Dr. Gaumer, who employed natives to collect, but they were 
necessarily from a limited area. My trip ended very much as it began, for I was 
obliged to return to Progreso, and from thence by steamer to Vera Cruz. 

BRITISH HONDURAS. 

British Honduras, or the colony of Belize, as it is often called, situated on the south- 
eastern shore of the peninsula of Yucatan, is about 160 miles from north to south and 
60 at the widest part. The navigation of the coast is both difficult and dangerous, 
on account of the numerous cays and coral reefs with which it is bordered. In the 
neighbourhood of the town of Belize, and for some distance inland, the ground is low 
and swampy and thickly clothed with mangroves and tropical jungle. Further west 
there is a narrow belt of alluvial soil, beyond which, and parallel to the coast, are 
tracts of arid sandy land called ' pine ridges,' from the pine trees with which they 



BRITISH HONDURAS. 25 

are covered. Still further inland are the ' Cahoon Ridges,' clothed with palm trees, 
while beyond are broad savannas studded with clumps of trees and intersected with 
streams. The Manatee Hills rise in a further succession of ridges parallel to the 
coast, and are from 800 to 1000 feet in height, while to the south the Coxcomb 
Mountains attain an altitude of 4000 feet; further inland there are said to be a 
succession of valleys and hills at altitudes varying from 1200 to over 3000 feet above 
sea-level, but this part is very imperfectly known. The climate near the coast is 
generally hot and damp, but tempered by the trade winds, and though the annual . 
rainfall is said to be about 100 inches, the country is tolerably healthy. Unlike the 
rest of Central America, British Honduras is not subject to earthquakes ; it appears 
to be entirely outside the volcanic area, which otherwise extends from Mexico to 
Western South America. 

In 1862 Salvin, as stated on p. 5, went from Coban by way of Peten, down the 
Belize River to the town of Belize on the coast, but unfortunately he left no details of 
the country through which he passed. His intention was to proceed direct to Yzabal, 
and thence back to the interior of Guatemala, but finding no vessel ready to sail, he 
hired a schooner and occupied the time in exploring some of the numerous atolls and 
coral-reefs which line the coast, and later published an interesting account of this 
expedition in 'The Ibis' for 1864. He described the Barrier Reef as extending 
from Ambergis Cay to Ranguana Cay, its most northerly point ; this last cay is twenty- 
five miles from the coast, so that the reef, instead of running more or less parallel with 
it, forms an angle enclosing u long lagoon, w^hich, as well as the reef, is studded with 
numerous cays. Nearly due east of the town of Belize, outside the Barrier Reef, and 
separated from it by a deep chaunel, lies the Atoll of Turneff, within which several 
lagoons are included. Fifteen miles eastward of Turneff lies another atoll, called 
Lighthouse Reef, on the eastern margin of which are four cays — Long Cay, Middle 
Cay, South- West Cay, and Soulh-West-of-All Cay j the remainder of the reef consists 
of a line of breakers, showing here and there a stranded log or a protruding spit of 
sand. It will be easily understood that these reefs, many of which are covered with 
mangroves and coco-nut palms, form an ideal place for sea-birds, and as Salvin's visit 
took place at the height of the breeding season, he procured a large number which he 
bad not previously obtained. 

In addition to these sea-birds, Salvin mentions two humming-birds, two tyrants, 
a warbler, a mocking-bird, an osprey, an ibis, egrets, etc. The paper quoted is too 
long to reprint in detail, but it is still the only account known to me describing 
the cays in question. His subsequent visit to the lagoons on the Pacific Coast of 
Guatemala in 1863 is referred to under the heading for that country. 



BIOL. CENTE.-AMER., lutrod. Vol., January 1915, £ 



26 PHYSICAL FEATUEES ETC. 



HONDURAS. 

The Eepublic of Honduras is bounded by the Bay of that name and the Caribbean 
Sea on the north, by Guatemala on the west, and Salvador, the Pacific Ocean, and 
Nicaragua on the south; it includes the islands of Ruatan, Bonaca (Guanaja), and 
the islands adjacent. The general aspect of the country is mountainous, and it is 
traversed by ranges and hills radiating from the base of the Cordillera. The main 
chain, which does not approach within 50 or 60 miles of the Pacific, is not an 
unbroken one, as it turns back and forms basins or valleys, within which are collected 
the head-waters of the streams which flow in the direction of the Atlantic Ocean. 
Viewed from the Pacific, the mountains present the appearance of a great natural 
wall, with a lower range bristling with volcanic peaks between it and the Western 
Ocean. The Cordillera proper forms an irregular line from north-west to south-east, 
interrupted, however, by the great transverse depression of Comayagua, which extends 
about 40 miles to the north with a width of from 5 to 15 miles, and contains the 
Humuya River, which discharges its waters into the Atlantic ; while to the south it 
forms the valley of the Goascoran River, which flows into the Pacific. The whole 
country has a great diversity of surface and elevation, with fertile valleys and high 
plains, afi'ording every variety of climate. 

Some notes on the aspect of this country were published in ' The Ibis ' for 1860 by 
G. Cavendish Taylor. He crossed from the Pacific to the Atlantic Coast, and many 
of the places described have been mentioned in the ' Aves ' section of this work. 
Arriving in December 1857, from Panama, at La Union, Salvador, he crossed to the island 
of Tigre, and thence to La Brea in the Gulf of Fonseca, which is studded with densely 
wooded volcanic islands. Tigre abounds with scorpions and large hairy spiders, and 
many birds were obtained near an old crater, now a lagoon full of reeds and floating 
grass. Mr. Taylor visited Aremecina, Caridad, San Juan, and Lamani, and so over the 
tolerably level plain mostly covered with forests to Comayagua. In the vicinity of 
the town were cactus bushes on one side, dense jungle intersected by rivers and woods 
on the other, and high mountains bounding the plain. Continuing his journey towards 
the Atlantic, Siquatepeque was reached by a route crossing the top of the mountains 
at an elevation of 5000 feet — here again was open plain, 3600 feet above sea-level, 
and surrounded by mountains ; then after passing over undulating ground covered 
with pine trees, giving it a park like appearance, he arrived at Taulevi. Here 
arrangements had to be made for the journey througli the dense forests to the Lake 
of Yojoa, some three leagues distant. Men had to be sent forward to clear a path 
and engage boats for the passage down the lake. The route lay at first by open 



HONDUEAS.-GtJATE.MALA. - 27 

savannas and wooded hollows, but soon a broad, rapid stream was crossed, and entrance 
was made into the dense forest, through which a road had to be cut, but further on 
the trees were lofty with huge buttresses at the base, and the undergrowth was not 
very thick. It was nearly dark when the river, flowing from the lake, was reached 
and the canoes launched. The current was slight, the water deep and still, and the 
banks were covered with high trees and dense forests, every bush and bough was alive 
with fire-flies, and the cries of night-hawks, coupled with the croaking of innumerable 
frogs, made no inconsiderable noise. The wind was ahead and occasionally so strong 
that the travellers were unable to proceed until it lulled, but at dawn they had 
completed twelve miles and had reached their destination. Two days were spent at 
Agua Azul, so called from the colour of the deep spring which rises near the 'hacienda' 
and flows into the lake. Numerous interesting birds were seen among the reeds and 
alligators [Crocodilus americanus) were not uncommon, while every tree and blade of 
grass swarmed with ' garrapatas.' Leaving the lake, which was surrounded by high 
mountains, the journey was made principally over savannas and open ground to the 
town of Yojoa, and via Potrerillos to the Atlantic. In the forest the route lay for 
miles through vistas of palm trees and bamboos, which shaded the path with their 
feathery branches, but unfortunately prevented the deep mud-holes from drying up. 
After leaving San Pedro, where brown monkeys with white faces were seen, a high 
range of mountains was crossed and Omoa reached on February 14th. 

G. C. Taylor enumerates one hundred species of birds, and G. M. Whitely sub- 
sequently sent us a collection of 135 species from the same country. These were 
named by Salvin and a list of them publislied in the ' Proceedings of the Zoological 
Society' for 1870, pp. 835-839. Whitely's skins (520) were obtained in 1869 in 
the vicinity of Puerto Caballo (Cortes) *, Julian, Medina, and San Pedro. The three 
last named places are situated in the low forest-lands on the Chamelicon Eiver ; San 
Pedro, the farthest inland, is not more than 30 miles from Puerto Caballo, now the 
Atlantic terminus of an incompleted inter-oceanic railway. 

After examining the collection carefully, it became apparent that the Ornithology 
of this part of Honduras scarcely difi"ers from that of the thoroughly explored lowlands 
of Vera Paz. 

GUATEMALA. 

Guatemala is coterminous on the north and west with Mexico, the flat low lying 
peninsula of Yucatan extends to the north-east, British Honduras, the Caribbean 
Sea, and the Republics of Honduras and Salvador are on the east and south-east, 
while on the south-west lies the Pacific Ocean. The greater part of the country is 

• Often confused with Puerto Cabello in Venezuela. 

b2 



28 PHYSICAL FEATURES ETC. 

mountainous, the main chain of the Cordillera forms the watershed, and at a mean 
elevation of 7000 feet runs nearly parallel to the Pacific coast at a distance of about 
fifty miles from it. The steep slope on the Pacific side is broken by many volcanoes, 
■while towards the Atlantic the land sinks in a gentle incline with subsidiary ranges 
extending nearly to the water's edge. Of the volcanoes several are active, the most 
noted is the Fuego (14,070 feet) with its twin sister the Agua, so called because 
in 1541 a lake, which occupied tlie centre of the crater, was discharged by a great 
eruption on to the former capital below and the city now called Ciudad Vieja was com- 
pletely destroyed by water. Both volcanoes are clothed with dense forests from about 
7500 feet to 10,000 feet, above which level there are scattered pines for 1000 feet 
or more, of which stunted examples are even to be found in the extinct crater of the 
Agua. Below 7000 feet the forest has been cleared for cultivation, and only parts 
are now clothed with a dense growth of scrub. The chief rivers are the Usumacinta, 
which flows into the Gulf of Mexico, and the Motagua and Polochic, which fall into 
the Bay of Honduras. Those flowing into the Pacific Ocean are short and rapid, as 
the fall from the Cordillera is very steep. 

The so-called ' coast country,' however, extends a long way inland, as during the 
wet season the torrents which descend from the Cordillera are charged with volcanic 
sand and disintegrated scoriae, and when discharged into the ocean they are cast 
back by the waves and the deposit forms a line of sand-bank. The constant heaping 
up of this bank often closes the mouths of the smaller streams during the dry season, 
and when the current is not sufficient to reduce the sand-bar the water expands inside 
the beach, forming lagoons and marshes along the whole coast. These lagoons are 
a favourite resort for waders and sea-birds, of which Salvin later obtained a large 
number, as well as a considerable quantity of fish. 

As previously mentioned, Salvin made no less than four expeditions to Guatemala, 
but I will only describe the physical aspect of the country we travelled over together, 
alluding, however, to those parts which he visited alone and giving extracts from 
some of his scattered papers published in ' The Ibis ' and other magazines. We 
landed in September 1861 at Yzabal, on the Golfo Dulce, and after a short stay we 
proceeded towards the Capital. The neighbourhood of Lake Yzabal is covered with 
dense forest extending beyond the Mico Mountains, which we crossed into the 
Motagua Valley in order to reach Quirigua. Here we spent a few days, and then 
proceeded up the river valley to Zacapa by mule path, the country gradually becoming 
drier and the vegetation more arid ; cacti and thorny shrubs became abundant, taking 
the place of more luxuriant plants. Indian settlements were found at intervals of 
every few miles, where the brushwood had been cleared for the cultivation of maize 
and coffee-trees, which were groAving in small patches. Much the same character of 
dry country prevailed throughout the journey to Guatemala City, which is situated on 



GUATEMALA. 29 

the Cordillera at an altitude of about 4500 feet above the sea. From the Capital we 
visited Dueiias, a village on the Pacific slope situated in a plain at the foot of the 
Volcau de Fuego and between it and the Volcan de Agua. Here we spent about 
three months, exploring the country and making frequent excursions into the forest 
of the Volcan Fuego, which furnished a great contrast to that near the coast ; the 
high trees were the only corresponding feature, but both vegetation and climate were 
entirely different. Instead of the incessant noise of the buzzing of myriads of insects, 
life seemed almost extinct and a dead silence reigned throughout, broken only by au 
occasional gust of wind or the fill of some rotten tree. The mountain itself is 
furrowed with deep ravines, called 'barrancas,' the sides of which are exceedingly steep 
and quite impassable, and in ascending the mountain, care must be taken to keep on 
the top of the ridges between them. The forest shuts out the view of the surrounding 
country, consequently landmarks are not available, and as one ravine almost exactly 
resembles another it is an extremely easy matter to lose one's way by inadvertently 
followins; the edge of a new ' barranca.' The lower part of the forest up to about 
7000 feet has been cleared for cultivation, but quickly reverts to a dense growth of 
scrub, above which is found a belt of evergreen-oaks followed by deciduous trees 
of various species, amongst which the remarkable Cheirostemon platanoides mingles 
in the highest range with alders. Then follows more open ground with pine trees 
and coarse grass, but the trees become stunted as the ascent increases, and finally 
disappear at an elevation of about 11,000 feet. From thence to the summit of the 
Fuego the cone is composed of cinders and ashes interspersed with short coarse grass. 
The mountain is divided into two peaks or cones near the summit, connected by 
a narrow ridge of cinders, the southern and higher peak is still active, and from it a 
perpetual column of thin smoke is always plainly visible. This cone is very steep, and 
the climb to the edge of the crater itself is exceedingly laborious, as the foot sinks at 
each step deep into the ashes. The view, however, from the point well repays the 
trouble of the ascent. Southward the eye travels a distance of 50 miles to the coast, 
far beyond again is seen in dim outline the horizon of the Pacific Ocean, while 
below on the other side lies the deep abyss of the crater itself. The northern cone 
is more or less covered with coarse grass extending to the summit, while the interior 
of the crater has been almost filled with the eruptions of the more recent southern 
volcano ; but signs of internal fires are not wanting, as jets of steam and sulphurous 
vapour are still seen issuing from the fissures in the rocks. In one of the hotter 
crevices 1 found a vigorous plant of Lycopodium clavatum and a Selaginella taking 
advantage of the warmth and moisture and growing with wonderful luxuriance at an 
altitude of nearly 14,000 feet. The descent was by no means easy, as there was no 
track to mark our way, but we had fortunately taken the precaution of slashing 
the trees with our big knives or ' machetes ' on our way up, which indicated our 



30 PHYSICAL FEATUEES ETC. 

route sufficiently to enable us to return to camp in safety. The Volcan de Agua is 
very similar to the Fuego, though somewhat less in height, but there is a fairly good 
mule tiack nearly to the summit. This track is frequented by the Indians, who 
ascend the mountain for the purpose of charcoal-burning and also in search of ice, 
which they found in the old crater in sufficient quantity to supply the needs of the 
Capital at that time. 

Having made considerable collections, we returned to Guatemala City and, recrossing 
the Motagua River and the Chuacus Eange of mountains, took up our abode at San 
Geronimo, a sugar-cane plantation in the plain of Salama, in Baja Vera Paz. The 
surrounding mountains are clothed with forest composed of various trees, including 
pines. The plain itself is arid, except when irrigated for cultivation, as at the 
Hacienda of San Geronimo. We next proceeded to Coban through the district north 
of the plain of Salama. The road soon leaves the plain, and the broken country is 
covered with scrub and forest, the rainfall being much greater as one approaches Coban 
than on the Pacific side of the Cordillera. During the rainy season there is usually 
a severe thunderstorm in the afternoon, followed by a clear sky, but during the dry 
season little or no rain falls and vegetation suffers greatly. In Alta Vera Paz and 
towards the Atlantic rain apparently falls at all seasons and all hours, and vegetation 
is consequently much more abundant. 

At Tactic, a forest district near the head of the Polochic Eiver, our porters failed 
to arrive, and we were forced to spend the night without our baggage. It was so 
bitterly cold that in the morning the ground and even the backs of our mules were 
covered with hoar-frost. A few days later, on our return journey, the effects of the 
unprecedently low temperature were plainly visible on the vegetation around. On 
reaching Coban Ave found a large Indian village where the inhabitants were born 
collectors, and very soon they brought in, in almost embarassing numbers, specimens of 
birds, frogs, toads, lizards, snakes, and insects of all kinds. The natives there were 
specially expert in the use of the blow-pipe, with which they killed most of the 
smaller birds. The weapon consisted of a straight piece of hollow wood about 
eight feet long, and the projectile, a hardened pellet of clay, fitted closely into the 
groove of the pipe and was blown from the mouth by the marksman. In this way 
a large number of birds was obtained with little or no damage to the plumage. Such 
was the accuracy of aim that, even at a distance of from 15 to 20 yards, many 
humming-birds were killed. 

After some weeks spent in collecting at Coban we visited Cvtbilguitz and Choctum 
in the low damp forest of Alta Vera Paz, thence travelling towards Salinas in the 
humid valley of the Chixoy or Rio de la Pasion, a tributary of the Usumacinta River. 
The roads or tracks made by the natives were extremely bad in this locality, the 
ground very broken, and the soil a stiff clay, so slippery in places that it was scarcely 



GUATEMALA. 31 

possible for animals to keep their footing. As there were no villages whence food 
could be obtained in this little known district, it was necessary to take a three weeks' 
supply from Coban and also to engage a number of Indians to act as porters. It 
was somewhat difficult to estimate the amount of food required per person, and for 
this purpose we decided to make a preliminary or trial trip extending over three 
days. We found that an Indian consumed daily about half his straw hat full of 
' topopoxti ' or baked Indian corn cake, and this with a few onions and ' frijoles ' or 
black beans supplied the necessities of life. Having arrived at the quantity required, 
we made up a sufficient number of loads and these were carried by the porters 
on their backs. 

I^n route we occasionally discovered a small Indian settlement, where our "mozos" 
found shelter in a hut formed of poles and thatched on the top and on two sides. 
These were resting places used by the natives on their way to Salinas in search of 
salt. Salvin and I preferred, however, sleeping in our hammocks slung to the trees 
in the adjoining forest, and as we were each provided with a waterproof sheet, we 
slept in dry beds notwithstanding the constant wet nights. 

The days were usually fine and were mostly spent in exploring the forest and 
collecting birds, insects, and plants. We remained a little over three weeks, till our 
supplies were exhausted, and then returned with our spoils to Coban. 

Owing to my having contracted an attack of fever and ague in the low ground 
at Salinas I was unable to accompany Salvin on his journey to Peten and Belize. 
On his second expedition to Guatemala, Salvin had already visited Lanquin and 
Cahabon, about three or four days' journey from Coban. He describes ('Ibis,' 
1861, pp. 138-149) the country as very wet and covered with forest, the roads 
— or rather tracks — impassable for animals, and all baggage had to be carried by 
Indians. The forests on the slopes of the limestone mountains were the home of 
the Quezal, the royal bird of the Aztecs, as well as of many other birds not found 
on the Pacific side, such as members of the families Cracida), Tinamidae, etc. Salvin 
says: "These forests are perhaps more worth seeing than anything in Guatemala, 
quite different to those on the West Coast, where the heat is excessive and mosquitoes 
and other insects abound and destroy one's comfort. In these forests it is otherwise; 
no ' garrapatas,' no mosquitoes, and a climate in the dry season which might challenge 
any in the world. Most parts are free from brushwood, and one may ramble where 
one pleases without being stopped by dense thicket. What strikes the eye most is 
the number of ferns, not only of plants, but of species ; every tree is clasped, every 
stone clothed with them, besides many of terrestrial habit." 

As soon as I had recovered from the effects of fever, I left Coban for Buenaventura 
on the Motagua River in order to collect fish, and the methods employed have already 
been described on a preceding page. 



32 PHYSICAL FEATUEES ETC. 

.Salvin then revisited his old quarters at San Geronimo, and taking his friend Robert 
Owen, the proprietor of the Hacienda, with him, he rode over the high land round 
Quiche and Totonicapam at an altitude of 10,000 feet. Here the climate is 
temperate, potatoes and wheat are largely grown, and on the uncultivated ground 
oaks, pines, and alders abound. Thence, crossing the Cordillera, he proceeded to 
Quezaltenango, a large town in the ' Altos,' and the capital of a considerable district, 
which he describes ('Ibis,' 1865, p. 187) as a corn growing and sheep producing 
hghland ; thence to Retalhuleu and on to the port of San Jose. At Retalhuleu he 
) ^»ard such glowing accounts of the prospect of obtaining a valuable collection of 
dea-birds and fish from the lagoons on the coast that he took a passage in a trading 
barque which was going from San Jose to Champerico to take in a cargo of coflFee and 
sugar, and succeeded in procuring a large number of specimens. 

When Salvin had finished collecting on the lagoons, he made an expedition to a 
belt of tropical forest parallel to the coast, but about twelve miles distant. Here it 
was that he specially remarked the contrast between the birds of the Pacific and 
Atlantic coasts — many of the most familiar birds of the low forest of Vera Paz, the 
Tiuamidse, Columha nigrirostris, and Ostinops montezuma, being entirely absent, nor 
does one find the genera Shamphocelus or Calliste, or the beautiful Cotinga amalilis. 
Much of the forest consists of bamboo, with here and there a huge tree standing high 
above it. Between this forest and the coast the soil is comparatively unproductive, 
bearing the stamp of land reclaimed from the ocean at no very distant date. The 
long line of volcanoes suggests a recent upheaval, and the constant discharge of sand 
by every river would tend to advance the coast by slow degrees. This low country is 
very subject to malarial fever — although Salvin escaped, his two attendants contracted 
it. Salvin returned to England soon after this, early in 1863, but ten years later, in 
the autumn of 1873, he paid his fourth and last visit to Guatemala, this time in 
company with his wife : although he added considerably to our collections the route 
taken was much the same as on previous expeditions. 

In this description of Guatemala, it must be remembered that when I visited it over 
fifty years ago there were no railways. There is now a railway from Puerto Barrios 
on the Atlantic Coast up the Motagua Valley to the capital and thence to the Pacific 
coast at San Jose, with a branch running from Mixtan near Escuintla to Eetalhuleu 
and Champerico. The country, therefore, is at the present time readily accessible by 
steamer from Belize and thence from the Atlantic port by train. 



SALYADOE.— XICAEAGIJA. 33 

SALVxVDOR. 

Salvador, though the smallest of the Central American Republics, is one of the 
most densely populated and largely cultivated, and consequently there is but little 
forest. It is bounded by Guatemala on the west and Honduras on the north and east. 
The country averages about 60 miles in breadth, and the coast line on the Pacific is 
160 miles in length. The seaboard consists of a comparatively narrow alluvial plain, 
beyond which is a plateau with a mean elevation of 2000 feet, broken by a number of 
volcanic cones lying to the south of the main Cordillera, and the whole Republic is 
very subject to earthquakes and volcanic outbreaks. The general aspect of Salvador 
led us to believe that the fauna and flora would be very similar to that of Guatemala 
and Honduras, consequently we neither visited the country ourselves nor did we 
employ any collectors there. 



NICARAGUA. 

The Republic of Nicaragua, wedged in between Honduras on the north and Costa 
Rica on the south, has a coast-line of about 2S0 miles on the Caribbean Sea and about 
200 on the Pacific. The land gradually decreases in width from north to south, while 
the main watershed extends eastward from within a few miles of the Pacific Ocean. 
Grey town (San Juan del Norte), at the mouth of the San Juan River, formerly 
possessed a fine harbour, but of late years the Colorado branch of the river, which 
bifurcates about twenty miles from the coast, now takes most of the water and the old 
channel and harbour have silted up. The main geographical feature of the country is 
the remarkable depression stretching for about 200 miles from the north-west to the 
south-east, parallel with the Pacific Coast and to the central plateau. This depression, 
which lies at a mean elevation of about 100 feet, is flooded by two great lakes, 
Managua and Nicaragua, which collect the drainage-water of the western provinces and 
also that from the eastern range of mountains, finally discharging it through the San 
Juan River into the Caribbean Sea, a distance of 120 miles. The Lake of Managua is 
about 50 miles in length and 25 in breadth ; the level is 16 feet higher than that of 
Lake Nicaragua, but the natural outlet, except in high flood, carries but little water, 
the surplus passing off by evaporation. The Lake of Nicaragua is about 100 feet above 
the sea-level and 150 miles long. Throughout its entire length this great depression 
is traversed by a remarkable chain of isolated volcanic cones, which, north of the lakes, 
takes the name of Marabios, terminating at the extreme north-west with Coseguina 
(4000 feet), and in the extreme south-east in the low wooded archipelago of Solenti- 
name and Chichicaste, near the entrance to the San Juan River. These volcanoes 
range from 4000 to over 6000 feet, while Momotombo, the highest point in the Republic 
of Nicaragua between the Gulf of Fonseca and Lake Managua, reaches an altitude of 

BIOL. CENTR.-AMER., Introd. Vol., January 1915. F 



34 PHYSICAL FEATUEES ETC. 

7000 feet. To the above mentioned series of volcanoes also belong those on the islands 
of Zapatero and Oinetepe, in the lake of Nicaragua. The latter, after a long repose, 
burst into renewed activity in 1883, and for seven days continued to spread devas- 
tation, destroying the crops and compelling the people to take refuge on the mainland. 
Several other volcanoes are still more or less active, and in 1835 Coseguina was the 
scene of one of the most tremendous eruptions on record. The outbreak lasted four 
days, during which time sand and ashes were carried to such a distance that they fell 
in Jamaica, Mexico, and Bogota. No rivers of any size flow westward to the Pacific, 
but the Lake of Nicaragua receives, near its outlet, the important Rio Frio from Costa 
Rica, which, at certain seasons, brings down a vast amount of water. Little is known 
of the region of rugged plateaux and savannas occupying fully half the country between 
the lacustrine depression and the Caribbean Sea. A large portion of the low ground 
is said to be covered with dense forest intersected by innumerable streams, all flowing 
eastward to the Mosquito Coast, which is low, swampy, and very unhealthy. 

Mr. Thomas Belt, a mining engineer and a well known naturalist, spent over four 
years at the gold mines of Santo Domingo in Nicaragua, and published an excellent 
account of his travels on his return in 1874. A considerable part of his book is 
occupied with extremely interesting observations on the Indians and the natural history 
of the district through which he passed, and I am indebted to him for the following 
details. Landing at Greytown he proceeded in an open boat up the San Juan River, 
which he describes as having a dangerous bar, over which he had to pass ; he then 
entered a wide channel with shallow water and beds of high grass on one side and a 
sandy shore on the other, in which alligators floated about like logs of wood and flocks of 
wading birds were seen in the marshes beyond. Proceeding up the river in still water, 
he emerged into a wider channel with a stronger current. The banks of the river 
were at first low and marshy, intersected by numerous streams fringed chiefly with 
palms and beds of wild cane and grass; further up the banks became higher and 
drier, and plantations of bananas and plantains were noticed in the clearings of the 
forest. About twenty miles above Greytown Mr. Belt reached the Colorado branch 
of the river, which now takes the greater part of the water from the lake to the sea 
by another outlet. There the banks were hidden by high trees laden with creeping 
and twining plants, many of which bore beautiful flowers, while beneath were tree- 
ferns with their light green foliage and slender stems. Higher up he passed the znouth 
of the Chiripo River, which rises in the interior of Costa Rica and joins the San Juan 
about thirty miles above Greytown. It is navigable for about twenty miles from this 
point, after which it becomes a rough mountain-torrent, and a mule track leads tlience 
to San Jose, the capital of Costa Rica. At Castillo, on the river bank, a considerable 
quantity of ' Ulli ' (ule) or rubber is collected by the Indians, which forms an 
important object of trade. This is obtained from a species of wild flg {Castilloa 
elastica), a plant with large leaves, diff'ering entirely from that found on the banks of 



NICAEAGUA. 35 

the Amazon [Siphonia elastica), and which is prepared in another manner ; the former 
is abundant in the forests of Nicaragua and Honduras. The San Juan River continues 
with much the same high forest on its banks as far as San Carlos, at the entrance to 
the Lake of Nicaragua; it is about 120 miles long and the lake is ]07 feet above the 
sea, so the water falls a little less than one foot a mile. The height of the lowest pass 
between the lake and the Pacific Ocean is said to be only 26 feet, and consequently 
this is the greatest depression in Central America between the Atlantic and Pacific. 
Owing to the enormous reservoir of water in tiie lakes, it has frequently been 
suggested as a practicable route for a ship canal between the two Oceans. 

On reaching the lake a sail was hoisted on board the little boat, and in a couple of 
days Belt arrived on the northern shore at Ubaldo, the landing-place for the machinery 
and goods destined for the mines at Santo Domingo. Leaving Ubaldo the road 
crosses some low rocky hills with scanty vegetation, consisting of spiny cncti, leathery 
leaved trees, thorny palms, prickly acacias, and bromelias with sharply serrated 
leaves; this being the dry season, the mule track was parched and dusty, though 
during the rains it becomes a slough of mud and water. The road led through the 
town of Acoyapo, which is in a grazing district with large cattle ' haciendas.' Soon 
after this. Belt crossed the range which divides the forest region extending from the 
mountains of Segovia to the Caribbean Sea, and separating it from the great lake 
depression. The savannas on this side were more humid and the moisture increased 
as he proceeded across the upper waters of the Mico River, which enters the sea 
at Blewfields, The black margin of a great forest, which had been visible for some 
time, was reached in the neighbourhood of Santo Domingo ; the ranges of irregular 
hills running mainly east and west were covered with vegetation, which was usually 
enveloped in a dense mist and produced a most depressing effect. The last part of the 
road was through brushwood, which had sprung up where the high forest had been 
cleared for planting maize ; but Belt soon found himself under a canopy of high trees 
the trunks of which were entwined with creeping aroids and lianas, sending down their 
great rope-like stems to the ground. This forest is always wet, and the undergrowth 
consists of small palms and magnificent tree ferns, witli thin stems and delicate foliage, 
and broad leaved heliconia;, leathery melastomse, and flesh-coloured begonias, with a 
variety of other damp forest loving plants. 

In 1872 Belt made a long journey to Segovia in order to engage labour, as the 
Indian miners mostly came from that province. The road lay over a rough forest 
country on the east side of the range dividing the great lake valley from Matagalpa, 
and this part of his journey strongly contrasted with any former one, as he was now 
at a long distance from the Atlantic, in a dry and arid region, due to the north-east 
trade wind having deposited its moisture on the intervening stretch of high land. 
Belt crossed several high ranges before reaching Ocotal, the capital of Segovia, situated 
near the sources of the Rio Wanks ; here grew pine trees and evergreen oaks at 

f2 



36 PHYSICAL FEATUEES ETC. 

])robably the southern limit of the former in Central America. Descending a steep 
slope beyond Ocotal, he came to a forest resembling that around Santo Domingo, 
though the trees were not so large, but tree ferns, palms, lianas, broad-leaved heli- 
coniae, and melastomse were again abundant, and he was told that the Quezal, the 
royal bird of the Aztecs, was occasionally met with. Belt, having successfully obtained 
the required number of Indians, returned by nearly the same route to Santo Domingo, 
and shortly after left for England. 

Mr. C. W. Richmond, who resided in Eastern Nicaragua from February 1892 to 
January 1893, when describing the climate, says [Proc. U.S. Nat. Mus. xvi. (1893)] 
that the east coast has a protracted rainy season of eight or nine months, with 
occasional spells of fair weather ; the rainfall is enormous, in some years reaching 
296 inches at Greytown. He went up the Rio Frio into Costa Rican territory, from 
the Lake of Nicaragua to the Guatusa Indian settlements, at the head of the canoe 
navigation. A shark, doubtless the same species as that known to inhabit the lake 
[Carcharias nicaraguensis), was seen as far up the river as he ascended. Later 
Mr. Richmond spent some months on the Escondido River, chiefly about 50 miles 
from the mouth. This river was formerly known as Bluefields, or Blewfields, and is 
probably the most important on the coast of Central America, with the exception 
of the San Juan. There is no troublesome bar, as is usually the case, and large ocean 
steamers ascend to Rama, 65 miles from the mouth, where two rivers, the Rama and 
the Sequia, join and form the Escondido. The banks for many miles, including both 
branches above Rama, are lined with banana plantations, the monotony of which is 
l)roken by the numerous picturesque ceiba and ebo trees which have been left standing 
in the clearings, while the dense tropical forest lies in the background. In the last 
15 or 20 miles of its course, the river winds through dreary silico swamps nearly 
devoid of bird life, and tlien empties itself into the Bluefields Lagoon, 15 miles 
long and 7 miles broad. Mr. Richmond enumerates 281 species of l)irds which 
he observed during this journey. 

COSTA RICA. 

This country was not visited by either Salvin or myself, and I am indebted to 
Mr. Carriker, Mr. Ridgway, and other writers for the following information. 

The little Republic of Costa Rica has an extreme length of 250 miles with a breadth 
of about 150. The greater part of the country is very mountainous, with narrow 
coastal plains on both sides, finally extending to about 30 miles in width in the north- 
eastern corner. The drainage system is complicated, the extreme northern portion is 
comparatively low, draining into Lake Nicaragua to tlie north and the San Juan 
River on the Atlantic ; while on the eastern side the streams and rivers have 
their sources in the high mountains, and descend rapidly through narrow valleys or 
gorges, separated by abrupt forest-clad ridges, which are frequently very narrow. The 



COSTA EICA. 37 

country is extensively wooded, the forests consisting of trees of all sizeSj many of them 
attaining enormous girth and height, especially on the eastern slope, where the rainfall 
is most abundant ; consequently, vegetation is there much denser and penetration very 
difficult. On the Pacific side the forests have less undergrowth, the trees are larger 
and taller, and progress through them is comparatively easy. There is, however, an 
exception in the Guanncaste region in the Nicoya peninsula, where great tracts of 
grass lands or savannas, with scattered patches of woodland prevail. Trees with 
berries and other kinds of fruit abound at all altitudes, furnishing food throughout 
the year for the multitudes of tanagers, finches, parrots, toucans, and trogons, while 
their blossoms give sustenance to innumerable humming-birds. The tree which, 
throughout the tropics, mostly attracts other birds is one of the Leguminosse, 
bearing biennially great masses of fragrant tassel-like blossoms which persist for some 
days before fading, while the season of flowering extends over a period of more 
than two months ; it is known as the ' guava ' (this is not, however, the guayava of 
commerce). 

On the Caribbean slope the rainfall is fairly continuous during the whole year, reach- 
ing on an average from 200 to 230 inches on the lower land. The gre;itest rainfall is 
from the middle of December to the middle of January, and again from the middle of 
June to the middle of August, Avhile from January loth to March 15th it is fairly dry 
and cool, and this season on the eastern slope is delightful. In the high regions the 
rainfall is less, and there are alternately six months of wet and dry season. During 
the first and last months of the rainy season the fall is slight and rarely of daily 
occurrence, while in July, August, and October it rains every afternoon, and often 
violently. During the dry season high winds prevail at all altitudes, but little if any 
wind blows during the wet season. Influenced by the constant rain, the vegetation is 
most luxuriant, and with it we find animal life consequently much more abundant. 
So numerous are the species that probably not less than three hundred and ninety 
land and fresh-water birds may be found on the lower portion of the Caribbean slope 
up to 3000 feet. The high peaks rising above the regions of the Central plateau 
constitute quite a distinct life zone. The main range of mountains extends from the 
extreme north-west to the eastern central portion, gradually increasing in height and 
ending in a chain of volcanoes reaching an altitude of from 8000 to 11,000 feet, 
including Poas, Barba, Irazu, and Turrialba. Here there is a break formed by two 
deep, broad valleys, the river lieventazon flowing in the one to the Atlantic and the 
Eio Grande de Tarcoles in the other to the Pacific ; their sources are within half a 
mile of each other, the ' divide ' or watershed being known as ' El Alto ' and having 
an elevation of 5000 feet. The whole country to the south is an unbroken mass of 
mountains containing but few inhabitants except the rapidly diminishing Indians. 

Mr. Carriker (Ann. Carnegie Museum, vi. 1910) considers that the Avifauna of 
Costa Rica is composed of three primal groups — the Boreal, the ISonoran, and the 



38 PHYSICAL FEATUEES ETC, 

Tropical; — the first two coming down frona the north, the last coming up from tlie 
south, and all meeting in the Republic of Costa Rica. He says, and truly, that 
the geographical position and meteorological peculiarities of the country make this 
enormous bird fauna within its confines explicable, but at the same time they greatly 
increase the difficulty of a satisfactory disposition of many of the species with respect 
to life zones. The continent decreases from 3000 miles in breadth to scarcely more 
than GO at the narrowest part of Costa Rica, and within those 60 miles are crowded a 
diversity of climatic conditions, altitudes, etc., scarcely paralleled in the Avorld. The 
northern and southern forms of the Pacific and Atlantic lowlands meet here and 
overlap, a single remnant of the Canadian bird-fauna persists on some of the isolated 
peaks of the high mountains (Junco vulcani), while modified forms of this northern 
species are found on the high lands of Mexico and Guatemala. A very large number 
of North American migrants arrive during the winter months and distribute themselves 
widely as to altitude throughout the country. Mr. Carriker also observes that amongst 
certain species there is a seasonal migration from a higher to a lower altitude, 
doubtless for the sake of food, and Salvin and I noticed similar migrations of 
several species of birds in Guatemala ; this was especially the case with humraing- 
])irds, which were very abundant at Dueiias during the flowering season, but in the dry 
season, when the flowers failed, this district was entirely deserted and the birds 
migrated to the Pacific Coast, where the atmosphere was damper and the blossoms on 
which they fed were abundant. 

The cultivation of bananas, however, is having a marked effect on the birds, which 
are decreasing at an alarming rate, for in the few years since the establishment of the 
Fruit Company at Port Limon the forest has been completely cleared — not only there, 
but also southward towards the Talamanca district. Very few of the forest birds 
frequent the banana plantations, and with the destruction of the forest they recede 
or disappear altogether. 

Mr. Ridgway (' Condor,' vii. 1905) gives an interesting account of Poas, which is 
the only active volcano in Costa Rica. After spending the night at San Pedro, 
he resumed his journey by bright moonlight at 4 o'clock the next morning. The 
' lecheria' or dairy farm at the upper edge of the cleared zone was reached soon after 
daybreak, and the primitive forest which covers the last thousand feet of the mountain 
was then entered. No pine trees were found here, as would have been the case at the 
same altitude in Mexico or Guatemala, as south of Nicaragua they cease to exist. 
The density of the forest was, however, sucli that it was impossible to leave the track 
without cutting a way with 'machetes,' and as the undei growth consisted mostly of 
slender climbing bamboos, with exceedingly hard stems, which almost filled the 
spaces between the trees, the difficulty of making much headway may be imagined. 
The variety of trees was very great, and all were laden with orchids, bromeliads, and 
mistletoes, the latter often conspicuously and brilliantly flowered and the bromeliads 



COSTA EICA. 39 

of dazzling hues of orange, scarlet, and crimson. Here, too, was seen the Quezal, amid 
surroundings no less magnificent than itself. 

Leaving the horses in the open basin of an ancient volcano surrounded by forest, 
Mr. Ridgway proceeded on foot to the summit of the cinder cone, but the view was 
disappointing, as the crater was filled with dense clouds, except for one moment, when 
a strong wind dispersed the mass of vapour and allowed a brief glimpse of the boiling 
lake, 400 metres below. From the summit a descent was made to the lagoon, another 
extinct crater filled with clear water almost icy in its coldness and surrounded by dense 
forest. The time spent on Poas was too short to learn much of the birds, but except 
at the summit they were everywhere found in great numbers. 

At Bonilla, an estate on the Atlantic slope near Turrialba, the roughness of the 
ground and the density of vegetation in the hot humid zone made it very difficult to 
collect, but in the cleared areas, where the ' potreros ' or pastures are found, the 
variety and abundance of birds was remarkable. In one locality at least four hundred 
species were obtained, and in the thick growth many escaped capture, while humming- 
birds were so plentiful that fifteen species were shot round one flowering ' guava ' tree. 
From here also was witnessed a flight of migrating hawks, passing northwards in 
hundreds and thousands. 

At Coliblanco, near Turrialba (6500 feet), the trees, including the brilliant scarlet- 
flowering Erythrina, were covered v.'ith creepers and epiphytes, while near the 
water-courses at least three species of magnificent tree-ferns were common, as well 
as Caladium-\\ke plants with leaves large enough to form a shelter against rain. 
As might be expected at this altitude, the birds Avere mostly difi'erent from those 
of Bonilla. 

Another interesting place was Pigres, at the mouth of the Rio Grande de Tarcoles, 
in the Gulf of Nicoya. Between this and the mainland proper lies the ' estero,' a 
broad creek of smooth water, bordered by dense mangrove swamps, somewhat narrow 
on the Pigres side where the land mostly consists of bare sand, but in places covered 
with trailing Ijwmosa, bearing broad leathery leaves and pink flowers ; matted clusters 
of thorny leguminous shrubs and thickets of low spreading mimosa-like trees inter- 
spersed with the poisonous manzanilla, formed the rest of the vegetation. Notwith- 
standing the dry season and the almost total absence of flowers, birds were very 
numerous in the vicinity of the village. Immediately beyond the fringe of mangrove 
swamps, on the other side of the ' estero,' a high mixed forest extended for many 
miles, the undergrowth consisting chiefly of small biscoyal palms bristling with 
long slender thorns of needle-like sharpness. Further inland these palms gave place 
to high cannas, and in the forest of tall trees, macaws, parrots, and parrakeets 
were numerous and noisy. 

An expedition from Coliblanco was made to the base of the cinder cone of Turrialba, 
about 9000 feet. Here everything was diff"erent from what had been seen below. 



40 PHYSICAL FEATUEES ETC. 

The several hundred acres composing the ' potrero ' resembled an immense well 
kept park, with long vistas through groves and clumps of magnificent trees on 
undulating grassy slopes, cropped closely by the grazing cattle. On the right rose the 
cone of the volcano, covered with dense ' chaparral,' or bush of evergreen oaks, 
while to the left the long ridge-like mass of Irazu was plainly visible. 

Mr. Ridgway made an ascent of this volcano, about 11,500 feet. The forest, of 
which but little remains, consists chiefly of oaks and differs widely from that of Poas 
and Turrialba. Higher up and close to the ash-cone the trees become scarce and 
scrubby, and finally only a growth of stunted Vaccinium-like shrubs exist. 

It is remarkable that in Costa Rica at least 700 species and subspecies of 
birds have been found. Dr. Outram Bangs, when alluding in the 'Auk' (1907) 
to the Costa Rican collections made by Mr. Underwood, remarks that the extensive 
bird-fauna of this small country, scarcely larger than the State of Florida, is due 
to the fact that the Central American forms extend to the Atlantic lowlands, while 
those from Panama and the south go up the Pacific slopes, separated only by the 
range of high mountains. 

An account of the Costa Rican Odonata, their larval forms and their habits, is 
given by Dr. Calvert in the ' Entomological News ' for July 1910. He and his wife 
remained in the vicinity of Cartago for a year, making collections of Odonata as well 
as of terrestrial molluscs, annelids, araneids, orthoptera, microdiptera, coleoptera, and 
lepidoptera to a smaller extent, but they were hurried away by the severe and frequent 
earthquakes which finally destroyed the town in May 1910. 



PANAMA. 

This State — or Republic, as it must now be called — comprises the neck of laud 
extending from Costa Rica to Colombia, an area equal in extent to about two-thirds 
the size of England and Wales, and forming the most southern country dealt with in 
the 'Biologia,' Very little, however, of the Isthmus of Darien, the land south-east of 
the Canal, has been visited by collectors. The main chain of the Cordillera decreases 
greatly in height towards the City of Panama, and between that place and Colon, where 
the railway and Canal traverse the country, the elevation falls to less than 300 feet. 
Salvin crossed by rail on more than one occasion, and spent some time collecting near 
the Station of Obispo, where he obtained a good many specimens ; but Enrique Arce 
and Mr. Champion were specially employed in Chiriqui, Arce subsequently proceeding 
to Veraguas where he remained for several years. 

The rivers, taken as a whole, are unimportant, but the Chagres with its tributary the 
Obispo attains formidable dimensions in the wet season, overflowing its banks and 
inundating a large area. 

The district immediately adjacent to the Canal has recently been described by Mr. A. 



PANAMA. 41 

Busck ('Report on the Mosquito Fauna of Panama,' 1908), who spent three months in 
the neighbourhood of Tabernilla near Colon. The ground slopes towards the Chagres 
River, and in the intervening country lies the bed of the old French sea-level canal 
which, even in the dry season, is covered with a series of shallow lakes connected by 
low marshes. Between these and the river are tall bamboos, sparsely interspersed with 
large hardwood trees, the crowns of which are covered with parasitic plants, orchids, 
and tillandsias, the last named affording a breeding place for several species of 
mosquito. 

When Mr. Champion visited Panama in 1881-1883 he investigated the Pacific slope 
only, that on the Atlantic side being very inaccessible, and except at Colon and along 
the railway, or near the coast, there were no villages or means of obtaining food or 
shelter. He endeavoured to ascend the Volcano of Chiriqui, which attains an elevation 
of 11,000 feet, but the only route through the forest lay by narrow tracks made by 
tapirs, and on reaching the summit of a ridge, at 8000 feet, further progress was im- 
possible, owing to the presence of an immense ravine, from which the upper part of the 
volcano could alone be seen. The night was spent in a hut erected by orchid collectors, 
but as no water could be obtained he was obliged to descend the following day. 

On the western slopes of the volcano the savannas reach an altitude of about 
6000 feet, while at 4000 feet cattle are pastured in large numbers for the Panama 
market. Higher up, to the north and west, a dense belt of forest covers the mountain 
side, but this does not extend to the summit. On the southern slope the forest had 
been cleared in many places for the cultivation of coffee, and a fine palm was locally 
abundant, but Coniferse were entirely wanting. At Chorcha (300 feet) the dense forests 
descended to the coast and interrupted the continuity of the large savannas bordering 
the Pacific Ocean. 

The Avifauna of Central America south of the Lake of Nicaragua, including Costa 
Rica, Chiriqui, Veragua, and Panama, is exceedingly rich. These countries contain 
more species than the whole of Europe, and nearly as many as the whole of America 
north of Mexico ; 432 species have been found in Veragua, including Chiriqui. 

Arce's collections, like those of other naturalists who have since visited the country, 
were almost, if not entirely, made on the southern or Pacific slope. 

Summarising his analysis of the birds of Veragua*, Salvin remarks (P. Z. S. 1870, 
pp. 178-179) as follows : — "The characteristic elenients of the Central American fauna 
consist not so much in the amount of generic peculiarity, which is very small, as in the 
fact that a very considerable portion of South American forms are here represented, 
not as specifically identical, but, in a large number of instances, as definably distinct 
in degrees of varying value. The element of the Central American bird-fauna to 
be traced to the northern continent, on the other hand, maintains a very different 
relationship to the bird-fauna of that continent. With the exception of a few species 

* Chiriqui was included by him under Veragua. 

BIOL. CENTR.-AMER., Introd. Vol., January 1915. o 



42 PHYSICAL FEATriiES ETC. 

isolated in the mountains of the higher portions of the Isthmus, and some others, we 
find that northern forms found in Central America are specifically identical with 
northern species, and that their presence is due in a great measure to migration during 
the winter season. As regards numbers, we find a gradual diminution as we recede 
from North America. These migrants are everywhere present, some few passing still 
further south into the equatorial provinces of the southern continent. Costa Rica and 
V'eragua, with Panama, possess these characteristics of the Central American fauna in 
the highest degree. It is here we find the greatest number of South American genera 
represented ; but the species are to a considerable extent not the same as the 
continental species. If we endeavour to account for the facts as we find them, by 
changes in past times in the physical features of the Isthmus, we seem to require : — 
1st. A union between Costa Eica, Veragua, and Panama with the southern continent, 
when those united lands possessed in common a much larger number of species 
specifically the same as at present, during which time the oceans may have been 
united north of Costa Rica. 2nd. The long duration of Costa Rica and Veragua 
as a ' continental ' island, when the union of the two oceans has been of greater 
extent. This period must be long enough to have established specific differences 
much as we now find them. 3rd. The emergence of the whole Isthmus in its present 
form. These requirements seem to fall in fairly with what has been demanded in 
other branches of natural science. Dr. Duncan requires a union in Miocene times 
between the oceans to account for the specific identity of certain corals. The union 
here demanded will suit my first and second requirements, I only regulate the amount ; 
and as for the period when it took place, fixing it to Miocene times would seem to 
answer to the requirements of the birds. That all the peculiar features of so varied 
a fauna can be accounted for by this theory I do not pretend to say. The changes in 
the physical features of the Isthmus, indicated by the numerous minor modifications 
of existing species, belong to the most recent events in geological history. To 
account for the greater differences observable we must go deeper into the abyss of 
geological time, where light is at present barely perceptible." 

In his first paper on this subject (P. Z. S. 1867, pp. 129-161), based upon less 
extensive material, Salvin stated that there was a closer affinity between the birds of 
Veragua and those of Costa Rica than between those of Veragua and of the Isthmus 
of Panama, but this proved not to be the case when Arce's later collections were 
examined. He then remarked that it was evident that Costa Rica and Panama had 
for a long period occupied the position of one or more islands between the two 
continents at a time when the two oceans were united by two or more channels ; and 
that an obvious division separating Costa Rica, Veragua, and Panama from the 
southern continent was a line drawn from the Atlantic Bay of San Bias to the mouth 
of the Bayano on the Pacific. 



PEARL ISLAjSTDS. " 43 

PEAEL ISLANDS. 

In the Bay of Panama there lies a small group of islands known as Las Perlas, 
or the Pearl Islands, consisting of Pedro Gonzales, San Jose, and San Miguel ; the 
last mentioned, also known as the Isla del Key, is by far. the largest, only twenty 
miles from the nearest mainland and sixty from Panama. This island is about fifteen 
miles long and irregularly oblong in shape, covered with low hills, which in turn are 
clothed with luxuriant tropical forest. The climate is hot and unhealthy, and the 
population consists almost entirely of negroes, who manage the affairs of the island 
and are very independent of the Panama Government. The pearl-diving industry 
having been almost abandoned, the people now grow vegetables, coco-nuts, and fruit 
for the Panama market. 

Mr. Champion, who visited San Miguel on our behalf in April 1883, was only there 
in the dry season, and the " luxuriant forest " of other writers he describes as 
" scrubby wood." The interior was somewhat inaccessible, the few paths or tracks 
leading only to the patches of cultivated ground. The coast, like that of the 
adjacent mainland, is covered with mangrove-swamps, which can only be traversed 
at low water. 

The Islands are in such close proximity to each other that probably the birds would 
be the same on each. On San Miguel 46 species were found, of which only four 
were considered by Mr. Bangs (' Auk,' xviii. pp. 24-33, 1901) to be well-marked 
island-forms, the remainder were, as might be expected from the semicircular form 
of the Coast of Panama, similar to those of the adjacent mainland. Some birds are 
undoubtedly carried across to the Islands by storms, but others perform the journey 
voluntarily, among them a small green humming-bird {Chloristilbon assimilis), which 
has been seen in perfectly calm weather flying straight for the Archipelago. 

Certain well known butterflies from the mainland also occur, including a Morpho 
(peleidesl), etc. 



2g 



[ 44 ] 



SOURCES FROM WHENCE OUR MATERIAL WAS OBTAINED. 



TuE following particulars as to the sources from whence our material was obtained 
supplement the account given in the Introduction, antea, pp. 11, 12. In Volumes I. 
of the ' Aves ' and of the ' Lepidoptera Ehopalocera ' the names of the various 
collectors who were specially employed to obtain material for the present work are 
recorded, viz. Messrs. F. B. Armstrong, E. Arce, G. C. Champion (1S79-1883), 
W. Lloyd, W. B. Richardson (1897-1898), H. Rogers (1877), and M. Trujillo (1888), 
and Mr. and Mrs. H. H. Smith (1888). From many others, also, we acquired or 
received vast numbers of specimens, all of which were duly acknowledged by our 
Contributors. 

As regards the Mammalia, Aves, Eeptilia and Batrachia, Pisces, Mollusca, 
Lepidoptera Rhopalocera, and Neuroptera (Odonata), the sources from whence our 
collections were derived are mentioned in detail in the * Introductions ' to the volumes 
on these subjects and need not be repeated. The Coleoptera, or beetles, however, 
numbering 18,029 species in all, and requiring eighteen volumes for their enume- 
ration, by twelve contributors, demand special notice. Notwithstanding their vast 
number, and the great abundance of individuals, this is the only group of insects, 
the Lepidoptera excepted, for which sufficient contributors have been forthcoming to 
enable us to complete their investigation. The Coleoptera, therefore, occupy in this 
work an undue amount of space amongst the thirty-eight completed volumes of 
Insecta, as compared with the Hymenoptera, Neuroptera, t&c, certain important 
families of which still remain unworked, this being due to the fact that the beetles 
have proved more attractive both to the collector and describer. 

The Coleoptera examined have been mainly supplied by the following col- 
lectors from each of the various countries : — Mexico — A. Salle, C. T. Hoge (who 
collected in nearly all the different States, Tamaulipas and Yucatan excepted), H. H. 
Smith (who specially visited Guerrero, Morelos, Vera Cruz, Tabasco, &c., for us), 
G. F. Gaumer (Yucatan), A. Forrer (N.W. Mexico, including the Tres Marias Is.), 
J. Flohr, F. D. Godman (Central and Southern Mexico), M. Trujillo (S. Mexico), 
Becker (Durango), Buchan-Hepburn (Chihuahua and Durango), and H. F. Wickham ; 
Bkitish Honduras — F. Blancaneaux ; Guatemala — A. Salle, O. Salvin, J. Rodriguez, 
G. C. Champion, L. Conradt, and F. Sarg; Nicaragua — T. Belt, E. M. Janson, and 
W. B. Richardson ; Costa Rica — H. Rogers, P. Biolley, Van Patten, C. F. Underwood, 



SOURCES or MATERIAL. 45 

and H. Lankester ; Panama (including Chiriqui, the Pearl Is., and Taboga) — G. C. 
Champion, E. Trotsch, H. Ribbe, and A. Boucard. The Hymenoptera examined 
and reported upon by P. Cameron mainly consisted of the collections made by 
G. C. Champion in Guatemala and Panama, and by G. F. Gaumer in Yucatan, most 
of the vast number of specimens obtained in Mexico by H. H. Smith having been 
received too late to be included ; it must be remembered, too, that the bees and social 
wasps have not been dealt vpith. The Lepidoptera, apart from the specimens captured 
by Salvin or myself, were obtained from the collectors who supplied us with the other 
insects; very extensive additions to both the Rhopalocera and Heterocera have, 
however, been made since 1900, especially by W. Schaus, who visited Mexico and 
Costa Rica in search of them. The Diptera examined were comparatively few in 
number, though H. H. Smith did his best in Mexico to make up the deficiency. The 
Rhynchota were very numerous, both in Heteroptera and Homoptera, as regards 
Guatemala and Panama, but till H. IT. Smith went to Mexico we had received 
very little from that country, the habitat of most of the previously described 
species from our region. The Orthoptera, again, were few in number, none of our 
collectors having paid very much attention to them. Tbe Arachnids were mainly 
supplied by H. H. Smith (Mexico), F. Sarg (Guatemala), and G. C. Champion 
(Panama) ; the Acaridea, however, were mainly procured by, and belonged to. 
Dr. Otto Stoll. In the Botany, the collections made by Mr. and Mrs. Salvin were 
used by Mr. Hemsley, but the study of the plants was almost entirely made from 
material contained in the Herbarium at Kew, including that which we had previously 
sent from Guatemala. Mr. Maudslay's work on the Archaeology was based on his own 
observations on the various ruins visited during his sojourn in the country ; Mr. J. S. 
Goodman, in his Appendix to this subject, gives an account of the system he used to 
decipher the Archaic Maya Inscriptions, which relate to a series of calendars. 

Mr. Champion's Itinerary is given on pp. 46-54, and the places he visited in 
Guatemala are easily traceable on Map 8 ; we are unable, however, to find space 
on our other maps to include all the Mexican localities quoted in the Zoology and 
Botany, some of which, indeed, cannot now be traced *. 

* It must be remembered that nearly all the names commencing with ' San,' ' Santo,' and ' Santa' have 
been used oyer and over again in tbe different States. The spelling of some of the others has been emended 
on our Maps to agree with official Mexican directories. 



[ 46 ] 



ITINERARY OF 
MR. G. C. CHAMPION'S TRAVELS IN CENTRAL AMERICA, 

1879-188 3. 



The only detailed itinerary supplied by any of our collectors is that of Mr. G. C. 
Champion, who, it must be remembered, devoted almost the whole of his time 
to entomology. A brief account of the numerous Guatemala and Panama localities 
he visited will be of interest to entomologists. His expedition to Central America 
extended from March 16th, 1879-May 23rd, 1883. The itinerary of his travels*, which 
is here reproduced with additional details, appeared in the American ' Entomological 
News' for February 1907, pp. 33-44. The Guatemala routes are shown on Map 8. 
During his stay in that country, March 16th, 1879-April 7th, 1881, he investigated 
both the Pacific and the Atlantic slopes, the central plateau, &c. From August 10th, 
1879-July 26th, 1880, he made his headquarters at San Geronimo, six miles from 
Salama, and made many distant expeditions to various parts of Alta and Baja Vera Paz 
from that place. The central plateau was traversed from Salama (via Rabinal, 
Cubulco, Joyabaj, and Quiche) to Quezaltenango, and also from the latter place to the 
Capital. The Pacific slope was explored from La Gavia to the Mexican frontier of 
Chiapas, at elevations between 1000 and 4000 feet. The ports of San Jose and 
Champerico, the lagoons at Paso Antonio, and the lakes of Amatitlan and Atitlan 
were visited. The Volcan de Agua was ascended on January loth, 1881. In Vera 
Paz, the Rio Dolores, near Cubilguitz, on the Coban-Peten road, was reached ; and 
the Polochic Valley was twice visited, once by boat from Panzos as far as the Lake of 
Yzabal. An expedition was also made across the Chuacus range of mountains from 
San Geronimo to Tocoy in the Motagua valley. Mr. Champion is of opinion that, 
from what he saw of the country during his travels, Guatemala might well be divided 
for zoological or botanical purposes into three (or f(jur) districts : — (1) " The Atlantic 
slope," which is mainly of limestone formaticm in Alta Vera Paz, and has a very long 
rainy season — in Cohan sometimes lasting into March, — and an abundance of humid 

* All made on horse- or male-back or on foot; the first Guatemalan railwa}-^ — from San Jose to Escuintla — 
was opened just before he left. 



ITINERAET. 47 

forest; (2) "The Central Plateau," including the 'Altos,' which is an arid upland 
region becoming higher towards the Mexican frontier and bordered southward by 
volcanoes which protrude from the main Cordillera — pines and oaks here clothe the 
mountain sides, on which cereals, maize, and, at high elevations, potatoes are culti- 
vated ; (3) " The Pacific slope," which has forest in the ' tierra caliente ' and on the 
mountain sides, now largely replaced by second growth (rastrojo) or cultivated with 
sugar-cane, cacao, or (at elevations up to 4500 feet) coffee. The plain of Salama, in Baja 
Vera -Paz, with San Geronimo at its eastern end — draining to the Atlantic, — is very 
hot and arid, abounding in Cacti, Yuccas, Agaves, &c., like the Motagua valley, from 
which it is separated by the Chuacus range of mountains. The mountains to the north 
of this plain, as shown by the abrupt change in the nature of the vegetation between 
Salama and Purula or Tactic, form the dividing line between the humid Atlantic slope 
and the plateau. These lower central valleys must either be included under district 2 
or treated as a separate faunal subregion. A comparison of the Lepidoptera Rhopalocera 
alone illustrates the great difference in the fauna of the two slopes * — many Erycinids, 
Ithomiids, Heliconius, Papilio, Leptalis, Thecla, &c., are peculiar to the Atlantic ; 
while Drucina, Euterpe, a few special Euptychia and Heliconius, a Morplio, &c., are 
found on the Pacific. The dry central plateau doubtless forms an impassable barrier 
for many species, and it has altogether a very restricted butterfly fauna. In the 
mountains in the neighbourhood of the plain of Salama a few peculiar Rhopalocera 
occur, such as Anxa nohilis and excellens, &c. The ' tierra fria ' or ' tierra helada ' 
(10,000 feet and upwards) produced no alpine or subalpine forms, merely stragglers 
from below. 

In Panama, April 17th, 1881-May 21st, 1883, Mr. Champion spent nearly all his 
time in Chiriqui, on the Pacific slope, between the Rio Chiriqui Viejo (near the Costa 
Rica frontier) and Tole, making his headquarters at the various coffee-plantations on 
the mountain-slopes, at an elevation of 3000-4000 feet, or else at David or Bugaba. 
The Volcan de Chiriqui was ascended on June 7th, 1882, to 8000 feet, and the 
Cordillera above Tole explored. The old route across the mountains from David and 
Caldera to the Chiriqui lagoon and the Bocas del Toro Islands, on the Atlantic side 
(used during the early days of the gold-mining in California, and before the Panama 
Railroad was finished), was found to be almost impracticable, and the northern slope 
was therefore not visited. The principal forests in Chiriqui are situated on the mountain 
slopes, in the low country to the east of David, and in the ' tierra caliente ' to the west of 
Bugaba and Divala; the forests alternate with extensive savannas along the lower part 
of the Pacific slope and in the country immediately adjacent to the western precipices 
of the Volcan de Chiriqui. The whole of the towns and villages are situated in the 
' tierra caliente,' and the Indians living in out of the way places in the Cordillera 

* Unfortunately this could not be very well shown in the Table of Distribution of the genera given in the 
Introduction to the Rhopalocera. 



48 



ITINEEAET. 



or on the northern slope are very seldom seen. On the southern slope of the Volcano, 
between 2000 and 4000 feet, a great deal of the forest had already been cleared (in 
1881) to plant coffee. San Miguel (Isla del R.ey) in the Pearl Islands and Tobago were 
visited by Mr. Champion, in April and May 1883, from Panama, and a certain number 
of insects, &c., were collected by him in these places. The absence of Conifers in the 
mountains, the paucity of Cacti, and the much less arid nature of the country afford 
a striking contrast to Guatemala, the fauna of Panama being very similar to that 
of Tropical South America. There is no arid central plateau in Panama, and the 
Cordillera, the loftiest part of which is in Chiriqui, decreases towards the isthmus, 
where it is only a few hundred feet high, so that the fauna of the two slopes is not 
likely to differ greatly. The Atlantic slope, however, has not yet been investigated. A 
Tenebrionid-beetle of the seashore, Phaleria dytiscoides, is recorded by Mr. Champion 
as common to the coasts of British Honduras on the Atlantic and of Guatemala 
and Nicaragua on the Pacific [cf. Coleopt. vol. iv. pt. 1, pp. 218, 219 (1886)], 
indicating a former connection of the two oceans at the isthmus of Panama. His 
itinerary was as follows * : — 



1879. March 16-18. San Jose de Guatemala, the Pacific 
port of arrival for travellers from Panama or San 
Pranciseo. Sea-coast, mangrove-swamps, lagoons, &c. 

March 19, 20. Travelling up to capital via Escuintla, 
by diligence, over execrable roads, all inches deep in 
dust at this (dry) season. 

March 21-ApriI 2. Guatemala city (about 4500 feet). 
Open plains, intersected by deep barrancas (ravines). 
Scrubby oak and pine woods in places on hill-sides. 
The volcanoes Pacaya, Agua, and Fuego visible to 
the southward. Many insects found on the banks 
of the streams in the barrancas. 

AprO 3-5. Ciudad Vieja. The first capital of Guate- 
mala, in the valley between the volcanoes Agua and 
Fuego. Coffee-plantations and cultivated ground, 
unsuitable for collecting-purposes. 

April 6-8. Guatemala city. 

April 9. Aceituno. Coffee-plantation near the capital. 

April 10-16. Guatemala city. 

April 17-May 12. Capetillo. Valley between the 
volcanoes Agua and Fuego. Coffee and sugar-cane 
plantations, with the forest-clad slopes of the Fuego 
adjacent. This estate is the property of the well- 
known Guatemalan naturalist, Juan J. Eodriguez, 
who has, from time to time, for upwards of thirty 
years, supplied the editors of this work with material 
from his district. 



1879. May 13-June 22. Zapote (about 2000 feet). On the 
forest-clad soutliern slope of the Volcan de Fuego. 
Broad, deep, dried-iip watercourses, full of great 
boulders, run downward through the forest here, 
making travelling diffiuult. The locality good for 
insects. Coffee cultivated. Some very fine forest 
passed through at San Cayetano, between Zapote 
and Capetillo, along the descending coast-road. 

June 24, 25. Antigua. 

June 26-July 21. Duenas (about 4500 feet). Near 
Capetillo. Coffee and Opuntia (for rearing the 
cochineal-insect) plantations adjacent to the Lake 
of Duenas. Various excursions made from here to 
Calderas, on the upper eastern slope of the Volcan de 
Fuego, up to about 7500 feet. Pines on the higher 
slopes. Earthquake-shocks frequent — the house be- 
longing to the owner of the estate (who had to leave 
it and live in Antigua) in ruins. The Acatenango 
peak of the Volcan, as well as the Brooking Fuego 
itself, conspicuous from Duenas. 

July 22-Aug. 7. Guatemala city. 

Aug. 8. Carrizal. Arid district with scrubby woods. 

Aug. 9. Llano Grande (about 2600 feet). Scrubby 
woods, cultivated ground, and pasture. Mule-trains 
and Indians bearing heavy loads constantly met with 
here, the route from the capital to Salama, Coban, 
&c., passing through this place. 



Elevations approximate only, taken from an uncorrected aneroid barometer. Probably too low in many cases. 



ITINEEARY. 



49 



1879. Aug. 10-Sept. 9. San Geronimo, Baja Vera Paz (about 
2950 feet). East end of plain of Salama, and six 
miles distant from the town of that name. Hot, 
dry region, with many cacti. Chuacus range of 
mountains adjacent, bordering the plain southward ; 1879. 
lower slopes clothed with pines, with forest of 
deciduous trees above. Sugar-cane and coffee planta- 
tions near village, belonging to English owners, the 
estate having a local reputation for the quality of 
the aguardiente (rum) and sugar ])roduced by them. 
Drainage to Atlantic. Headquarters for about one 
)'ear. Many long excursions made from here to 
distant places in both Alta and Baja Vera Paz, on 
the Atlantic slope. Some fine butterflies (Ancea spp.) 
peculiar to the Chuacus range. Various Longicorn 
(Ochresi7ies),JiupTesiid(Acma'odera),and other genera 
of Coleoptera characteristic of the drier portions of 
Mexico occur on the plain of Salama. A large 
Buprestid {ChalcopJiora vin/intensis) in the pines on 
the Chuacus slopes. An Ithomiid butterfly (Dircetvin 
klugi) seen swarming in the shady garden of the 
hacienda. 

Sept. 10. Santa Barbara (about 4450 feet). Moun- 
tainous region east of San Geronimo. Scattered 
woods. Pines below. Cultivation of maize, &c. 

Sept. 11, 12. Santa Cruz (5500 feet). Mountainous 
region of Chilasco, the watershed between the 
Motagua and the Polochic, east of Santa Barbara. 
Scattered woods of Liquidambar, &c. Forest appa- 
rently all cleared in viciuity of village. Nights very 
cold here. 

Sept. 13-Oct. 1. San Geronimo. 

Oct. 2-5. Purula (about 4000 feet). Open ground 
with a humid virgin forest adjacent on mountains. 
A new track through the forest towards Cerro Verde, 
an excellent entomological locality, the road to Sabo 
also productive. These localities again visited in 
April or May, 1880. The 'quezal' {Pliaromacrus 
modnno) not rare in the dense forest on the Cerro 
Verde road, and a ' Howler ' {Mycetes villosus) 
frequently heard between Purula and Sabo. 

Oct. 6, 7. Sabo (2900 feet). Clearing made in dense 
humid forest to plant coffee, on a steep mountain- 
slope. A very productive locality. Pulex irritans, 
however, swarmed to such an extent in the disused 1880. 
hut used for sleeping-quarters that it was impossible 
to remain very long in the place. 

Oct. 8-15. Panima * (1800 feet). Hot, narrow valley 
of the Rio Sinanja, a tributary of the Polochic. 
Valley, mostly cultivated with maize, &c., followed 

• Misprinted ' Paneina ' on many of the labels attached to insects in the 
collection made by Mr. Champion, and therefore wrongly quoted in some 
of ibe volumes of this work. 

BIOL. CENTR.-AMER., Introd. Vol., January 1915. 



down to near Rihaco and upward towards Matanza. 
Road from Sabo descending very abruptly through 
forest in which a transparent-winged Pieriue-butterfly 
(DismorpJiia foriunata) was abundant. 

Oct. 16. Purula. 

Oct. 17. Cachil. Open arid mountain-slopes north of 
the plain of Salama, with agaves, palms {Thrinax), 
&c. A peculiar Euptijchia (ruhricata) taken here. 

Oct. 18-Nov. 3. San Geronimo. 

Nov. 4-6. Tocoy (about 2000 feet). Arid district on 
the Zacapa road. 

Nov. 7. El Jicaro, near the Village of Guacamaya, on 
eastern slope of Chuacus rauge. Scrubby woods, 
pines above. 

Nov. 8-12. San Geronimo. 

Nov. 13, 14. Purula. 

Nov. 15. San Miguel Tucuru (about 2000 feet). 
Polochic valley. Mostly cultivated ground, cotton, 
maize, &c. 

Nov. 16. La Tinta. Polochic valley. Tropical vege- 
tation. Indigo formerly cultivated here, hence the 
name. 

Nov. 17-23. Senahu (2800 feet). Limestone moun- 
tains north of the Polochic valley. Humid forests, 
cleared in many places for coffee-plantations. Long 
rainy season, but water scarce, rapidly disappearing 
underground. A district rich in land-shells. Again 
visited in June, 1880. 

Nov. 24-30. San Juan (1800 feet). A small coffee- 
estate, no village, on the mountain-slopes north of 
the Polochic. Forest mostly cleared. Mountains 
of the Republic of Honduras visible to the S.E. 

Dec. 1, 2. La Tinta. 

Dec. 3-7. Tamahu, a few miles higher up the valley 
than Tucuru (about 2250 feet). Mostly cultivated 
ground. 

Dec. 8. Santa Rosa (about 4000 feet). 

Dec. 9-28. San Geronimo. 

Dec. 29. Tactic (4300 feet). Scattered Liquidambar 
and other deciduous trees. Forest all cleared to near 
the inaccessible mountain-tops. Large Indian popu- 
lation in district, hence the continuous clearing of 
the trees to plant maize (' milpas '). 

Dec. 30, 31. 1 Coban, Alta Vera Paz (about 3800 feet*). 

Jan. 1, 2. ) Humid region, rainy season sometimes 
extending into February or March. Forest nearly all 
cleared to plant coffee, maize, &c. Numerous German 
traders and planters settled here. The residence for 
many years of a keen zoologist, F. Sarg. Large Indian 
population. 

Jan. 3. Tactic. 

Jan. 4-6. San Geronimo. 

* Maudslay makes it 4280 feet. 
H 



50 



ITINERA ET. 



1830. Jan. 7. Buenaventura. 

•fan. 8-12. Gualeinalii city. 

Jan. 3 3. Lake of Arnatitlan (about 3450 feet). Arid 
district, with many ' nopales ' (plantations of Opuntin 
for rearing the cochineal insect, all enclosed within 
dusty adobe walls). The Volcan de Pacaya not very 
far distant. 
Jan. 14-18. Guatemala city. 
Jan. 19. Buenaventura. 
Jan. 20-25. San Geronimo. 
Jan. 26. Santa Rosa. 

Jan. 27-Feb. 1. San Joaquin, Alta Yera Paz (about 

3200 feet). Pine-clad, arid mountain-slopes. Rio 

Chisoy (or Chixoy) below. About the northern limit 

of the arid region of the central plateau. 

Feb. 2. San Cristobal (42.50 feet). Cultivated ground 

adjacent to the Lake of San Cristobal. 
Feb. 3-5. Balheu (Valeu) (3850 feet). Pine-clad 

mountain-slopes. 
Feb. 6-10. Cohan. 
Feb. 11. Chiacam (2400 feet). Coffee-plantations and 

scrubby woods. 
Feb. 12. Siin Agustin Lanquin (1000 feet). Lime- 
stone formation. Second-growth woods ; forest all 
cleared. Bio Cahabon, a large tributary of the 
Polochie, adjacent. An unproductive locality. 
Feb. 13-23. Cahabon (Cajabon) (about 800 feet). 
Second-growth woods, forest all cleared near village. 
Large Indian population. No other people here, 
except the priest (who kindly accommodated stray 
travellers) and two or three Guatemalan officials. 
The very large church here is placed on a hill, higher 
than the smaller ones on which the Indians have 
built their huts. Plantations of cacao, maize, &c., 
around the village. Toucans seen in these places. 
Feb. 24-28. Lanquin. The large limestone cave 
visited*, in company with an Austrian plant-collector 
casually met in the village. Only insects seen in the 
cave, an apterous Orthopteron {Arachnomimus 
cavieola). The llio Cahabon, a broad stream, issues 
from its mouth, making its first appearance above 
ground at this place. 
Feb. 29-March 6. Chiacam. 
March 7-9. Coban. 

March 10-20. Cubilguitz (1050 feet), near Choctum. 
Limestone region, with humid forest on the hills, the 
roads between the hills extremely bad. Broad vallevs 
with scattered trees. The Rio Dolores reached, but 
not crossed. On main road from Cob.an to Peten, 
the track to Salinas (a place where salt is obtained 
and sent on Indians' backs to Coban) turning off to 

* A\to entered by Sahin, on March 8tl), 1860 [cf. ' Ibis,' iii. pp. 140, 141 
(18«1)]. 



A good locality, but 
from the scattered 



the westward near Cubilguitz. 
food absolutely unobtainable 
Indian residents. 
1880. March 21. Satchicha (2000 feet). In fording the 
river here my mule was so badly cut about the legs 
by the jagged knife-edged submerged limestone 
ledges that it was unlit for work for three mouths 
afterwards. 
March 22-24. Coban. 
March 25. Tactic. 
March 26-April 13. San Gerdnimo. 
April 14-23. Purula. 
April 24-28. Panima. 
April 29-May 2. Sabo. 
May 3. San Miguel Tucuru. 

May 4-12. Cliaeoj, near Chamiquin, sometimes called 

La Uamaca (from the old suspended rope-bridge over 

the Rio Polochie) (about 500 feet). Tropical forest, 

with many palms (mainly Attalea cohune), which 

decrease in size as the Polochie is ascended. A 

new iron bridge in course of construction (in 1880). 

A very good entomological locality. 

May 13-18. Teleman, on the Rio Polochie, the upper 

limit of navigation for small boats from the Lake 

of Yzabal. Tropical forests, with many palms, the 

leaves of the lofty Atlalea cohune arching across the 

road. Culicidse swarn)ing. Unhealthy district. The 

' Howler ' frequently seen in the trees near the river. 

Some peculiar Syntomid - moths found amongst the 

prickly herbage. 

May 19-22. Panzos, on the Rio Polochie. Tropical 

forests, &c., as at Teleman. "Willows on river-bank. 

Unhealthy district. My first attack of fever here. 

May 23. Danta, on northern shore of the Lake of 

Tzabal, about fourteen hours' journey by small boat 

from Panzos. Many alligators and ' Howlers ' seen 

on my way down, manatees also noticed in the lake. 

Culicidae swarming. 

May 24-25. Travelling up the Rio Polochie, two days, 

against stream. 
May 2o-June 2. Panzos. Culicidse (especially a sooty- 
black species) so bad here, even by day, that it was 
almost impossible to do more than a few hours' 
collecting at a time. Local name for them, 
' zancudos.' 
June 3-14. Senahu, travelling up from Panzos by way 
of Trece Aguas. [The American entomologists, 
Messrs. H. S. Barber and E. A. Schwarz (of the U.S. 
National Museum at AV^ashington) have visited this 
district during recent years, and they succeeded in 
obtaining various minute Coleoptera of the same 
species captured by myself in 1880.] 
June 15-21. San Juan. 



ITINEEART. 



51 



1880. June 22, 23. Chacoj. 

June 24. San Miguel Tucuru. 

June 25-27. Purula. , 

June 28-July 26. San Geronimo. My last (8th) visit 
to this place. 

July 27. Eabinal (2850 feet), en route for the ' Altos.' 
Dry region, with scrubby woods, cacti, agaves, yuccas, 
&c., as on the plain of Salama. 

July 28. Cubulco (2900 feet). Similar country. 

July 29. Joyabaj (4300 feet), an Indian village, reached 
by a long precipitous ascent from Cubulco. Open 
mountainous region, intercepted by deep barrancas. 
Pines and oaks on slopes. Central plateau becoming 
higher westward. An interesting butterfly {Chri/so- 
phanus pi/rrhias) seen in numbers on the way up from 
Cubulco. 

July 30. Santo Tomas Chiche (6100 feet). Los Altos 
region. 

July 31-Aug. 5. Chimente, Quiche Mountains (7600 
feet). Indian village. Oaks, pine, alder, &c., on 
slopes. Potatoes and maize cultivated. Cyanide 
bottles (with the results of a day's collecting) stolen 
from saddle-bags here. 

Aug. 6. Totonicapam (7900 feet). Pine-clad slopes, 
but too far from the town for collecting purposes. 

Aug. 7. Chevuc (9900 feet). Pine woods. Be- 
nighted at this place, having missed the road along 
the Cordillera to Los Encuentros in the dark. 

Aug. 8. Los Encuentros (8400 feet). A resting- 
place for the night, on the road to Solola or the 
capital. Arid open ground. [Conradt collected 
insects at Tecpan, a place to the eastward.] 

Aug. 9, 10. Descousuelo (Solchicha) (about 10,500 
feet). Pine-forests. Very bleak situation above 
Totonicapam. Carriage-road from Quezalteuaiigo to 
Guatemala city passes this place, following the 
summit of the highest portion of the Cordillera. 
Potatoes only cultivated. A very good locality *, 
most of the Coleoptera, and some of the Lepidoptera 
Heterocera, collected proving to be new, but no 
peculiar butterflies were met with. 

Aug. 11-15. Pachoc (or Patch oc) (9200 feet). Indian 
village. Pine-woods. Only accommodation obtainable 
a small schoolroom used during the day, the bare 
earth covered with pine-branches serving as a bed. 
Slopes of the Cordillera accessible from this place. 

Aug. 16. Totonicapam. 

Aug. 17-19. Quezaltenango (7600 feet). Cultivated 
ground mostly. Large Indian population. Un- 
productive locality. 

* Tlie in§eots from Degconsuelo and Pachoc were colleclively labelled 
'Totonicapam' (a town at the foot of tbe Cordillera) in the collections 
made by Mr, Cliampion. 



1880. Aug. 20-Sept. 9. Einca of Las Nubes on the southern 
(Pacific) slope of the Cerro (or Yolcau) Zunil, abovfj 
Mazatenango (4050 feet). Extensive coffee-planta- 
tions, with dense forest above. The most producti\e 
locality visited on the Pacific slope. Several new 
butterflies (species of Dnwina or Euptijclna) met with. 
An interesting bird {Oreophasis derbiaaus), a monkt-y 
(Ateles ater'!), &c., seen in the forest. An enormous 
Passalid-beetle {Proculus goryi) found commoiilv 
beneath the large tree trunks left on the ground to 
decay in the ' cafetales ' (coffee-i)lautations). 

Sept. 10-23. San Isidro (1600 feet). 'Tierra caliente,' 
Pacific slope, below Mazatenango. Second-growth 
woods, cleared in places for coffee and cacao planta- 
tions. A Cnlujo plentiful in the tangled under- 
growth, but difficult to secure. 

Sept. 24. Retalhuleu (950 feet). Similar country. 
Now connected by rail with the port of Champerico. 

Sept. 25-Oct. 7. Las Mercedes (3200 feet). Pacifio 
slope. Immense coffee-plantations in this Costa 
Cuca district. Kearly all the original forest clcareil. 

Oct. 8-17. El Reposo (800 feet). Low country near 
Pacific. Mostly second-growth woods, but sonje 
forest, with lofty palms, in vicinity. Macaws 
(guacamayo's) often seen. 

Oct. 18, 19. Paraiso (300 feet). Near Pacific. Scrubby 
woods, bamboos, &c. 

Oct. 20-22. Champerico. Sea-coast. The remarkable 
fish (Anahleps dovii) seen in the lagoons. [Also met 
with by Salvin, at Chiapam, near here.] 

Oct. 23. El Reposo. 

Oct. 24-26. Las Mercedes. 

Oct. 27, 28. Coatepeque (1250 feet). Mostly culti- 
vated ground. [Village almost completely destroyed 
by earthquakes during recent years.] 

Oct. 29. Rio Naranjo (450 leet). Second-growth 
woods. Mexican frontier adjacent. A new iron 
bridge in course of construction. 

Oct. 30. Near Naranjo. Benighted by losing road. 

Oct. 31-Nov. 3. Finca La Union (22.50 feet). El 
Tumbador district, department of San Marcos. 
Coffee-plantations and second-growth woods. No 
villages hereabouts, the town of San Marcos on 
higher ground in the Cordillera. 

Nov. 4-7. Finca La Carolina (2600 feet). Tumbador 
district, overlooking lower portion of Soconusco, 
Chiapas, to Mexican coast, the Volcanoes Tacana 
and Tajuniuleo visible to the north-west. Unpro- 
ductive locality. 

Nov. 8. La Union. 

Nov. 9. Bio Naranjo. 

Nov. 10. Coatepeque. 

Nov. 11-14. Las Mercedes. 

U2 



52 



ITINEEART. 



1880. Nov. 15. San Martin, near Oistuncalco (7400 feet). 
Scattered trees, ground mostly cultivated. 

Nov. 16-18. Quezaltenango. Cerro Qnemado visited. 

Nov. 19-Dec. 14. Las Nubes (Cerro Zunil). [The 
village of Santa Maria, on way down from Quez-al- 
tenango, almost destroyed during recent years by an 
eruption of the volcano of that name.] 

Dec. 15. San Isidro. 

Dec. 16-26. San Agustin (2250 feet). Southern 
(Pacilic) slope of the Volcan de Atitlan. Second- 
growth woods, coffee-plantations, Ac, all very dry 
and dusty at this season. Many Lepidoptera 
Heterocera taken ' at light ' in the verandah of the 
house. 

Dec. 27-29. San Lucas Tollman (4900 feet). Indian 
village on the borders of the Lake of Atitlan, which 
is very deep and has no visible outlet to the Pacific. 
Onk-woods, &c., arid region. Pampojilaj, on the 
coast road, visited, and the lake crossed (at night) to 
northern side. Numerous thickly populated Indian 
villages round the lake, the Indians not very friendly. 

Dec. 30. Panajachel (4900 feet). Stayed at a flour- 
uiill, close to tlie lake. The town of Solola not very 
far distant. 

Dec. 31. San Lucas Tollman. Returned from Pana- 
jachel by a tortuous detour along the high ground 
above tlie lake. 
1881. Jan. 1. Godines (6900 feet). Arid district above the 
precipitous cliffs bordering the Lake of Atitlan on 
the N.E. side. Magnificent view at sunset of the 
lake and the adjaceut volcanoes and mountains to the 
southward during the cloudless skies of the dry 
season. 

Jan. 2. Chimaltenango (5650 feet). Reached by way 
of Patzun and Patzity.ia. Upland district cultivated 
with cereals. 

Jan. 3-11. Guatemala city. 

Jan. 12. Antigua. 

Jan. 13. Yoloan de Agua (about 13,000 feet) ascended 
at night (moonliglit) from the Indian village of Santa 
Maria (6500 feet). ]5elt of deciduous trees above 
the cultivated ground to about 9500 feet, scattered 
pines above, even in crater. Path up deep and 
extremely narrow, between dense tussocks of high 
grass. Very few insects met with on summit, the 
butterflies seen merely common stragglers from 
below. Indians ascend to fetch a little ice obtained 
from holes made in the ground. So windy that it 
was impossible to remain long on summit, clouds 
forming rapidly after about 10 a.m. 

Jan. 14. Antigua. 

Jan. lo-Feb. 3. Pantaleon (1700 feet), Pacific slope. 
' Tierra caliente^' Sugar-cane fields and second 
growth (rastrqjo). 



1881. Feb. 4-14. Mirandilla (1700 feet). Similar ground. 
Feb. 15. Escuintla. 
Feb. 16-18. Torola (1000 feet). Scattered patches of 

forest. 
Feb. 19-28. Paso Antonio (400 feet). Open savannas, 
with scattered Crescentia anil other trees, near Pacific, 
above Istapa. I^agoons here adjacent to the Rio 
Michotoya drained and used for pasturing the hungry 
cattle in dry season. Silurians and other fish 
captured in large numbers by the natives when tlie 
water becomes low in the lagoons, and iguanas also 
sought after, for food. Culicidoe and ticks very 
troublesome. Some new Dytiscida; and other aquatic 
insects taken. 
March 1-3. Torola. 

March 4. Savana Grande, near the Rio Maria Linda 

(about 1150 feet). Scrubby woods, pasturage, and 

cultivated ground. Passed through village of 

Guanagazapa. 

March 5. La Gavita (La Gavia) (1700 feet). Similar 

ground. 
March 6, 7. Brito. 

March -8-16. Torola. District swarming with ticks 
in dry season, and extremely unproductive, entomo- 
logically, like the rest of the 'tierra caliente' of the 
Pacific slope, during this period. 
March 17-April 3. Guatemala city. 
April 4, 5. Escuintla. 
April 6, 7. San Jose de Guatemala. Left by coasting 

steamer on 7th en route for Panama. 
April 9. La Union, Salvador. Landed for a few 

hours. 
April 10. Corinto, Nicaragua. Landed for a few 

hours. 
April 12. Punta Arenas, Costa Riea. Landed for a 

few hours. 
April 17-20. Panama city. Left on 20th by small 
steamer for Chiiiqui ; the " port " reached by a 
tortuous passage through mangrove-swamps, navi- 
gable at high-water. 
April 22-30. David. Principal town of Chiriqiii. 
Open savannas, with scattered, leathery-leaved, deci- 
duous trees. Denser growth by river - side and 
■on hills adjacent, the latter productive, entomo- 
logically. 
May 1-27. Finca Nance Bonito (about 2800 feet). 
Southern slope of the Volcan de Chiriqui. Plenty 
of forest, cleared in places tor coffee-plantations, a 
fine palm locally abundant beneath the larger forest- 
trees, at about 2000 feet, just above the limit of the 
savannas. Coniferse altogether wanting in the district, 
probably not reaching south of Nicaragua. Two 
species of monkeys {Cehics hypohucus and an AUlex, 
almos-t certainly geoffroyi) seen. Productive locality. 



ITLNEKAET. 



53 



1881. May 28-June 25. Las Potrerillas (Bl Banco) (about 

'2500 feet). Similar ground, not far from Nance 
Bonito. 

June 26-July 13. David. 

July 14-Aug. 1. Pinca La Elvira. Similar to Xance 
Bonito. 

Aug. 2-8. Las Potrerillas. 

Aug. 9-11. Boquete( 3550 feet). South-eastern slope 
of the Volcan de Chiriqui. 

Aug. 12-20. La Caldera( 1500 feet). Savanna region, 
with scattered trees. On old route from David to 
Atlantic coast, practical on foot only, but rarely 
used, all provisions having to be carried. 

Aug. 21-Sept. 27. Las Potrerillas. 

Sept. 2S-0ct. 20. David. 

Oct. 21-Nov. 28. Bugaba (1000 feet). Fine forests 
here *, extending into Costa Eica, eastward and south- 
ward savannas, with scattered trees. Eio Ascaria and 
Eio Chiriqui Viejo descending through forest-comitry. 
) Sugar-cane and coffee cultivated to a small extent. 

The place of residence of the brothers Trotsch and 
other collectors formerly employed by Staudinger, 
the well-known Lepidopterist. 

Nov. 29. Jugales. Eesting-place on way up to the 
Potrero del Volcan. 

Nov. 30-Dec. 10. Potrero del Volcan (4000 feet). 
Eoad from about 1000 feet above Bugaba rapiilly 
ascending through dense forest. Savannas up to 
(5000 feet, to the precipitous western slope of the 
Volcan de Chiriqui (about 11,000 feet), and to the 
abruptly descending banks of the Eio Chiriqui Viejo. 
Dense forests westward and southward, in wliich 
peccary are not rare, and jaguars, &c., occur. Large, 
numbers of cattle for the Panama market pastured on 
these savannas. Monkeys {Ateles sp.) and peccary 
{Dicotyles labiatus) sometimes shot for food by the 
' vaqueros.' Uninhabited district. 

Dee. 11, 12. Bugaba. 

Dec. 13-26. David. 

Dec. 27-31. 

1882. Jan. 1-Feb. 2. 
Feb. 3-5. Camaron (1750 feet). Milpas (maize-fields) 

in forest-clearings, on Costa Eica road. 

Feb. C-March 15. Bugaba. 

March 16-21. Las Potrerillas. Country in a smoky 
haze at this season, due to the general burning of the 
undergro«th. Fires dangerous to tiie thatched 
houses, owing to the strong ' northers.' 

March 22-April 10. Bugaba. 

April 11-13. David. 

April 14-18. Bugaba. 

April 19-29. Potrero del Volcan. Second visit, more 

• Probably all destroyed by this time. 



[ Bugaba. 



• productive tiian first. Many large Lamellicorn- 
beetles (Fliisiotis, various Dynastids, &c.) found 
floating in the water of the large shallow lagoons in 
the forest, on the banks of which numerous trees 
frequented by the insects were in flower. The short- 
tailed ' quezal ' {Pharomacrus costcn-icetisis) seen at 
about 5500 feet. 
1882. April 30-May 30. Bugaba. A very good locality at 
this season, the commencement of the rains. New 
clearings in forest — made to plant rice, maize, or 
tobacco — productive in Coleoptera, Ehynchota, &c. 
A large number of interesting Lepidoptera Ehopalo- 
cera occur in the district, including Morpho cifpris 
and other species of the genus. A marmoset, Chryso- 
thri.v oerstedi, occasionally seen in numbers on the 
outer limits of the forest ; also an occasional peccary 
{Dicotyles tapaju). 

May 30-June 5. Las Potrerillas. 

June 6-8. Slope of the Volcan de Chiriqui (5500 feet). 
Staying in hut erected by Swiss orchid collectors on 
the very steep forest-dad slope. Ascended to 8000 
feet, by tracks made by tapirs through the dense 
undergrowth of bamboo, &c., to summit of ridge. 
Further progress stopped by an immense precipitous 
ravine, beyond which the upper part of the volcano 
could be clearly seen through the trees. The vege- 
tation on the summit consisted apparently of low 
bushes (? Vaccinium). No water could be found near 
hut and a longer stay not possible. Some interesting 
new Lampyridse, Hispidae, and other Coleoptera found 
in these places. 

June 9-12. Las Potrerillas. 

June 13, 14. Finca Nance Bonito. 

June 15- July 3. Bugaba. 

July 4. Jugales. 

July 5-7. Potrero del Volcan. 

July 18-Aug. 15. Bugaba. 

Aug. 16-19. David. 

Aug. 20-28. Bugaba. 

Aug. 29, 30. Divala (Vivala) (350 feet). Savannas, 
covered in places with a short prickly Mimosa, the 
Eio Chiriqui Viejo adjacent. Tropical forest, with 
many lofty palms, westward, A ' road ' into Costa 
Eica passes this place, not many miles from the 
Pacific. Eastward, on the savanna, is the old 
capital of the district, Alanje. Abandoned saw-mill 
noticed on one of the streams. 

Aug. 31. Mosque (700 feet). 

Sept. 1-Nov. 30. Bugaba. 

Dec. 1-3. David. 

Dee. 4-27. Bugaba. Garrapatas (ticks) swarming in 
the bush diu-ing dry season, necessitating a frequent 
change of clothing. Boas occasionally met with. 
Insects scarce, except near water, butterflies mostly 



54 



ITINERARY. 



■nom. Many species of latter found about llie 
scattered pools of the nearly dry Eio Ascaria at this 
seasou, mostly Nymphalids and Hesperiids. Culieida) 
not very troublesome. 

1882. Dec. 28-30. David. 

Dec. 31. Chorcha (300 feet). Dense forest, descending 
down to the mangrove-swamps of the coast, inter- 
rupting the continuity of the large savannas border- 
ing the Pacific east of David, said to be infested with 
snakes. 

1883. Jan. 1. San Lorenzo (200 feet). Savannas with woods 

here and there. 

Jan. 2, 3. Los Remedies. A small port, reached 
through mangrove-swamps. Savannas inland. 

Jan. 4-10. Tole (1150 feet). Similar country. Ex- 
cursions made from here to various places in Cordil- 
lera. Cattle-breeding the principal business here. 

Jan. 11. Cerro Algodon (2000 feet). 

Jan. 12-17. Pefia Blanca (3000 feet). Very broken 
open country in vicinity of the Peiia (Blufl"). Slopes 
of the mountains covered with forest. Ascended to 
about 5500 feet. Indians from distant places on 
the Atlantic (northern) slope assemble in an unin- 
habited spot in the Cordillera near here annually. 
Dancing in the open air is kept up for two or three 
days, till most of the men and women become hope- 
lessly drunk from the aguardiente supplied by 
travelling traders. The adults seen at one of these 
functions were tattooed with blue and red pigment. 

Jan. 18. Cerro Algodon. 

Jan. 19-22. Tole. Nearly the whole of my money 
stolen from saddle-bags at this place, a return to 
David therefore imperative. Savannas east of Tole, 
on the Santiago de Veraguas road, visited, but found 
to be unproductive at this (dry) season. Several 
interesting Malacoderm-beetles {Astylue, Lycostomus, 
<Stc.) were, however, taken from flowers. Very little 
cultivation seen. 

Jan. 23-28. Nancito (800 feet). 

Jan. 29. Los Bemedios. 



1883. Jan. SO-Feb. 3. San Feliz (650 feet). Savannas, \\\i\x 
patches of wood. 

Feb. 4, 5. La Isleta. Adjacent to the Itio Foiiseca. A 
'Howler' {Alycetes palliatus) seen in the trees along 
the streams. 

Feb. G, 7. Cana Fistula. Near the Montaiia do 
Chorcha. 

Feb. 8-10. David. 

Feb. 11-March 3. Bugaba. 

March 4-9. David. 

March 10, 11. Bugaba. 

March 12, 13. David. 

March 14. Boquita, on way down to coast. 

March 15-17. Boca Chica, near coast. 

March 18-24. En route to Panama in small coasting, 
vessel carrying various passengers, who were accom- 
modated in the open air on planks above the cargo 
of pigs. Punta Mala passed with difficulty. 

March 25, 2C. Panama. 

March 27. Colon (Aspinwall). Many butterflies seen 
from train on way across isthmus, but nest to none 
found at Colon. 

March 28-April 2. Panama. 

April 3-27. San Miguel, King Island (Isia del Rev), 
Pearl Islands. Scrubby woods much cleared in places 
to plant yams, &c. Tangled jungle, with some large 
trees, in southern portion of island. Coast fringed 
with mangroves and coco palms. Darien coast and 
mountains visible from San Miguel. A few interest- 
ing insects obtained. Unhealiliy place. Too unwell 
to do much collecting. IS'o horses or cattle. Tracks 
available along beach at low water. 

April 28, 29. On way back to Panama in small boat. 

April 30. Panama. 

May 1-17. Taboga Island, Bay of Panama. Rocky 
ground, ascending to about 800 feet, with small 
streams. Pine-apples grown on the slopes, often 
stolen by sailors. Many butterflies and other insects 
occur on the island. 

May 18-21. Panama. -».* 



[ 55 ] 



LIST OF COMPLETED VOLUMES. 
Zoology, Botanv, and Archaeology. 



The 21-5 Parts of Zoology, 25 of Botany, and 17 of Archaeology are divided into 
63 Volumes, of which a complete list, with their contents, is given in tabular form 
on pp. 85, 86. The analysis of the contents of each of these volumes supplies the 
names of the contributors and other particulars, and, in the case of the Zoology, 
a brief summary of the author s views on the nature of the Fauna, as stated in 
their Introduction. The 215* Parts of Zoology form 52 volumes — one devoted 
to Mammalia, four to Aves, one to Reptilia and Batrachia, one to Pisces, one to 
Mollusca, four to Arachnida, one to Chilopoda and Diplopoda, and thirty-eight to 
Insecta. The Crustacea, Protobracheata, Vermidea, &c., have not been studied, 
mainly for lack of material. Amongst the Insecta, too, no worker has been found for 
certain groups of Hyraenoptera, Diptera, Rhynchota-Homoptera, and Neuroptera, and 
these omissions are specially noted in the analysis of the volumes dealing with the 
Orders in question. The 25 Parts of Botany form five volumes — four of text and one 
of plates. The 17 Parts of Archaeology form four volumes of text, together binding 
into one volume, with an additional common titlepage, four volumes of folio plates, 
and an Appendix (text only), the whole subject thus extending to six volumes — two 
of text (quarto) and four of plates (folio), 

* Part 211 was issued in two sections : " A " in Dec. 1911, " B " in May 1912. 
Part 212 „ „ „ : " A" in Tab. 1913, " B " m Aprill914. 



[ 56 ] 



ANALYSIS OF CONTENTS OF EACH VOLUME. 



1. The 'Introductory Volume' includes the general preface to the whole work, 
Zoology, Botany, and Archajology, a complete list of the subjects contained in each 
Volume, general statistics, maps, &c. The ' Contents ' of this Volume, which forms 
the first of the entire series, are given in detail on p. iii. 

ZOOLOGY. 
2. Mammalia : by E. R. Alston, with an Introduction by Dr. P. L. Sclater. 

The author of this Volume died on March 7th, 1881, before the enumeration of the 
species was cfinduded. The MSS., however, of the Supplement (pp. 203-212) was 
left by Mr. Alston in an almost complete state, and was finished by Mr. O. Thomas in 

1881. The Tables (I.-VIII), printed in the Introduction, were drawn up at our 
request by the author shortly before his death, and Dr. P. L. Sclater gives an 
analysis of them in his Introductory remarks on the subject (pp. x-xix), published in 

1882. The total number of species enumerated is 181, of which a complete list is 
given in the Introduction (pp. iv-ix). These fall into five categories : (i.) Nearctic, 48 ; 
(ii.) Neotropical, 65 ; (iii.) Neogean, 17 ; (iv.) Autochthonous, 47 ; and (v.) Intro- 
duced, 4 (Mures). The eight Tables drawn up by the author show : I. General dis- 
tribution; II. Nearctic species (17) found in the Northern States of Mexico, but not 
recorded from south of 25° N. lat. ; III. Nearctic species (18) found in Central and 
South Mexico, but not recorded from south of the Isthmus of Tehuantepec ; 
IV. Nearctic species (8) extending to Guatemala and Honduras, but not recorded 
from south of Nicaragua ; V. Nearctic species (5) extending to Costa Rica and 
Panama, but not recorded from south of the Isthmus of Panama ; VI. Species (17) 
common to the Nearctic and Neotropical Regions ; VII. Species (47) peculiar to 
Central America, or not yet ascertained to occur elsewhere ; VIII. Distribution of the 
Neotropical genera. Dr. Sclater (p. xiv) summarizes the analysis of the Central 
American Mammal fauna as follows : — " It may fairly be said that (excluding the 
introduced Mures) at least 100 of the constituent species are essentially Neotropicalin 
their character or have Neotropical affinities, while of the remainder not above 60 can 
be said to be decidedly Nearctic. There is therefore no doubt that the Central 
American isthmus, at any rate as far north as Tehuantepec, should be assigned to 
the Neotropical Region, of which it should be deemed to constitute a distinct province, 
characterized (1) by the incursion of a considerable number of Nearctic forms, 
especially in the northern districts, (2) by the presence of a certain number of peculiar 



ZOOLOGY. 57 

species of Neotropical genera, and (3) as being the focus of the families Procyonidse 
and Geomyidaj, two well marked groups of Mammals which have extended alike 
into the Nearctic and Neotropical Regions." 

The twenty-two coloured plates include figures of thirty-four species, a list of which 
is given on p. xx. 

3-6. AvES: by O. Salvin and F. D. Godman : Vols. I.-III. (text), III. completed 
with the assistance of Dr. R. Bowdler Sharpe and Mr. Ogilvie-Grant ; 
IV. (plates). 
Three Volumes are required for the enumeration and description of the 1413 species 
of Aves belonging to the Central American fauna, and a fourth for the 84 plates. 
Vol. I., published in 1879-1887, gives an account of a portion of the Passeres, the 
families Turdida; to Alaudidae ; and, on the conclusion of the work, in 1904, an 
Introduction to the whole subject was issued, with Tables (pp. xi-xxxviii) showing the 
geographical distribution of the families and species represented in Mexico and Central 
America. Vol. II., published in 1888-1897*, includes the continuation of the Passeres 
and the whole of the Macrochires, Pici, Coccyges, and Psittaci. Vol. III., published 
in 1897-1904, includes the Striges, Accipitres, Steganopodes, Ilerodiones, Phcenicopteri, 
Anseres, Columbae, Gallinic, Geranomorphse, Limicolae, Gavise, Tubinares, Pygopodes, 
Alc6e, and Crypturi. Vol. IV. contains the whole of the Plates and a complete list of 
the 149 species figured. Salvin's long continued ill health, and sudden death in 1 898, 
retarded the conclusion of the Third Volume, and this was subsequently finished with 
the assistance of Dr. Sharpe and Mr. Ogilvie-Grant. For this reason, too, all idea of 
a Supplement was abandoned, notwithstanding the large amount of additional material 
which had come to hand during the progress of the work. The additions, however, 
were mainly amongst the Passeres, completed in 1892. The Introduction to Vol. I. 
(1904) contains an account of the authors' various expeditions to Central America, 
the sources from whence their material was obtained, &c., and the following par- 
ticulars as to the nature of the Bird-fauna of the region : — To summarize the results, 
the Avifauna of Central America may be described as essentially Neotropical, with 
certain peculiar forms restricted to it. The fifteen families represented are all rich in 
endemic forms, and the families themselves are almost all tropical. On the other 
hand, a large number of species belonging to the more widely distributed genera find 
their winter home in Mexico and Central America, or further south, returning to 
breed in the Nearctic Region, even Humming-birds and others wandering far north at 
this season. The data is insufficient to show the lines of migration of all the species. 
Some, no doubt, travel southward from the United States to the mainland of Soutli 
America by way of the Caribbean Sea or the West Indian Islands, perhaps just 

• The permanent Titlepage and ' Contents ' were issued in 1904. 

BIOL. CENTR.-AiiER., Introd. Vol., January 1915. 1 



58 CONTENTS OF EACH VOLUME. 

touching the eastern portion of our region en route ; others probably find their way 
down to the central tablelands, and a few western species, again, pass over the low 
lands of the Pacific coast. 

Nearly half (636) of the 1413 species enumerated are treated as endemic, fifteen 
of the families — Trochilidne, Fringillidae, Tanagridse, Formicariidae, Dendrocolaptidae, 
Troglodytidae, Turdidoe, Psittacidaj, Phasianidae, Peristeridije, Cotingida?, Trogonidae, 
Cracidoe, Tinamidse, and Rhamphastidse — having many peculiar forms. Compared 
with America north of Mexico, the fauna of the region here dealt with is particularly 
rich, the numbers being. North America (1S95) 768 species as against Central 
America 1413 ; while India (with Ceylon and Burma), with nearly double the area, has 
(1898) only 1626 species. All, or nearly all, the new species were described first in 
the ' Ibis • or « P. Z. S.' by Salvin. 

Birds, from their power of flight and the habit of migration common to a large 
number of them, are much more easily distributed than most other vertebrates, and, 
therefore, do not throw the same light on the subject of geographical distributif)n 
as in the case of more sedentary animals. This must, nevertheless, be applied in 
a general sense, for many of the species are extraordinarily local. 

In dealing with the Aves the limits of the region have been extended to include 
the llevillagigedo Islands on the Pacific side, on account of the numerous sea-birds 
inhabiting them ; the Island of Old Providence on the Atlantic side, which has 
a humming-bird peculiar to it ; and some places on the Isthmus of Darien. 

The eighty-four coloured plates illustrate 149 species. 

7. REPTiLuand BATEAcnrA: by Dr. A. Giinther. 

The author, in his Introduction published in 1902, summarizes his remarks on 
geographical distribution as follows: — The general features of the Reptilian and 
Amphibian Faunas of the area under investigation have been satisfactorily ascertained. 
Forming the connecting link between the two Neogean regions. Central America 
possesses a Reptilian and Batrachian Fauna with the various constituent elements so 
mixed that, if only certain families or genera were taken into consideration, almost 
every district of this area could be associated with either the North- or South-American 
region. The tropical Fauna, as we proceed from lower to higher latitudes, gradually 
changes or is replaced by that of the temperate region ; but this change is not 
uniform throughout the breadth of the land, and the two faunas frequently overlap 
in deep and manifold indentations. Tropical types are found to preponderate in 
the low lands of the Atlantic side, which expand into the broad Yucatan peninsula, 
and on the humid slopes of moderate elevation ; some extend to, and even reach 
northwards of, the Rio Grande. On the Western side tliey are found in similar 
localities, but in a narrower belt, along the Pacific coast. On the other hand. 



ZOOLOOxT. G9 

numerous types of the southern North American Fauna are spread over Northern 
Mexico, extending along the Central American plateau to the extreme limits of our 
area, and even beyond. This southward extension of northern types is due partly 
to the identical physical conditions of the arid tableland of Sonora and Chihuahua, 
which is merely a continuation of that of Arizona and New Mexico, and partly to the 
great altitude and temperate climate of the Central American plateau. Thus, a 
boundary line between the North and South American regions cannot be draA\n: 
Central America forms a transition-tract unlike any other part of the world, showing 
the most extraordinary diversity of climatic, physical, and meteoric conditions 
within comparatively small areas, favouring the evolution of a great variety of types 
of genera and species, and influencing the dispersal of immigrants from the North 
and South. 

The range of the 695 species enumerated is shown in the Table appended to the 
Introduction (pp. x-xvii). 

Since the conclusion of Dr. Giinther's work, Dr. Hans Gadovv has twice visited 
Mexico (1902 and 1904), mainly to study the distribution, &c., of the Amphibians and 
Reptiles. He collected specimens of 135 species. The following papers have been 
written by him : — 

(1) " Evolution of the Colour- pattern and Orthogenetic Variation in certain 

Mexican Species of Lizards, with Adaptation to their Surroundings " [Proc. 
Eoy. Soc. Lond. Ixxii. pp. 109-125, pis. iii.-v. (1903)]. 

(2) "The Mexican Axolotl" ['Nature,' Ixvii. pp. 330-332 (1903)]. 

(3) "The Distribution of the Mexican Amphibians and Reptiles" [P. Z. S. 1905, 

ii. pp. 191-245]. 

(4) " A Contribution to the Study of Evolution based upon the Mexican Species of 

Cnemidophorus" (P. Z. S. 1906, i. pp. 277-375, pi. xx.). 

(5) 'Through Southern Mexico.' London, 1908. 

(6) " Geographical Distribution of Animals " [' Darwin and Modern Science,' pp. 319- 

336 (1909)]. 

(7) " The Eflfect of Altitude upon the Distribution of Mexican Amphibians and 

Reptiles" [Jahrb. Jena Abt. f. Syst. xxix. 1910, pp. 689-714]. 

He also published an excellent account of the portions of Mexico visited during his 
travels, and we give elsewhere a copious extract from one of his papers. 

During recent years attention has been called to various Reptilia, Batrachians, 
Arachnids, Coleoptera, tScc, living in the Bromeliads on the branches of trees. 
Specimens of Spelerpes variegatus and Hyla godmani were obtained from these plants 
by one of our collectors, Mateo Trujillo, in Mexico, and otlier species have since been 
captured by Dr. Gadow. 

I 2 



60 CONTENTS OP EACH VOLUME. 

8, Pisces : by C. T. Regan. 

The Introductory remarks to this subject (published in 1908) are arranged by the 
author under five headings : — (1) Principal Faunal Works on the Fresh-water Fishes 
of Mexico and Central America; (2) Principal Collections described in this Work ; 
(3) Classification ; (4) Geographical Distribution, illustrated by two separate maps ; 
(5) The Shore-Fishes of the Atlantic and Pacific Coasts of Mexico and Central 
America. The geographical distribution is discussed at length under the headings to 
the separate families, ten of which are represented in the region, that of the Cichlidse, 
Percidse, Characinidaj, Catostominae, and Cyprininse being illustrated by shaded maps in 
the text. The author considers that the Nearctic and Neotropical Regions are quite 
distinct, and to explain the distribu^tion of fresh-water fishes he says that the volcanic 
chain of mountains which stretches across Mexico from Coliraa nearly to Vera Cruz 
may be taken as the boundary. The Nearctic Region he divides into three subregions, 
one of which, the " Lerma," lies wholly within Mexican territory. The Neotropical 
Region is also divided into three subregions, the Central-American subregion again 
being separated into four " provinces," Balsas, Guatemalan, San Juan, and Isthmian ; 
the limits of these are shown on Map 2. 

Mr. Regan says that the Cichlidae, which form a very large family of fresh-water 
fishes, is the dominant perciform group in Tropical America and Africa. In America 
the Cichlid?e extend from Texas to Argentina, and comprise about 150 species. Africa 
appears to be somewhat richer in both genera and species, whilst three representatives 
occur in India and Ceylon. A map is given in the text to show this distribution. He 
states that the Mexican and Central American Cichlids must have originated in the 
Southern Continent. 

The Percidaj are fresh-water fish inhabiting Europe, Northern Asia, North America 
east of the Rocky Mountains, and Northern Mexico. Fossil Percidae are found in 
the Eocene deposits of Wyoming. 

The distribution of the Characinidae, also fresh-water fish, is equally interesting. 
They extend from Northern Mexico over all the South -American continent, except 
the extreme south, and are found also in Africa, but not in Madagascar. 

These two families and the Lepidosirenidse support the theory that a land 
connection between South America and Africa may have persisted until the beginning 
of the Tertiary period. 

The Catostominae extend from Guatemala all over North America and into Eastern 
Asia. 

The Cyprininse are found from Canada southward to the Lerma Valley in Mexico 
and throughout Africa and Eurasia, except in the extreme north. They abound 
in Borneo, but are absent from Celebes. 

Amongst the Shore-Fishes entering fresh water, eighteen Pacific coast species are 
specially noted as having a closely related representative on the Atlantic. 



ZOOLOGY. 61 

Mr. Began concludes his remarks on these fishes as follows : — " The fact that in so 
many cases species may be paired is more in harmony with the view that there has 
been a gradual modification during isolation than with the supposition that a mutant 
has arisen which has replaced the parent form." 

The twenty-six plates include figures of seventy-eight of the 415 species enumerated. 
The Maps inserted in the Introduction showing the distribution of certain families are 
worthy of attention. 

9. Terrestrial and Fluviatile Mollusca : by E. v. Martens. 

In this volume the author summarizes the characters and distribution of the species 
in common Tables, instead of giving separate descriptions. The geographical distri- 
bution is given at great length in the ' Introduction ' (published in 1901), accompanied 
by separate Tables of the Terrestrial and Freshwater genera. The difference between 
the fauna of the Pacific and Atlantic slopes is noticed, but is stated to be not very 
important as regards the land shells. Central Guatemala (Baja Vera Paz), however, 
is said to be occupied chiefly by metamorphic rocks, while North Guatemala (Alta 
Vera Paz) is mostly of limestone formation : this diff"erence accounts for the greater 
richness of land shells in the latter province. One peculiarity of the fauna of the 
Pacific slope is worthy of note, viz. the occurrence of large sized Otostomi and BuUmuli 
in Western Mexico related to various Andean forms, the distribution being somewhat 
analogous to that of the majority of the Cactacese. The submarine Mollusca of the 
eastern and western shores are said to be distinct, more so than some truly marine 
shells, but there are some remarkably analogous forms among them. 

Of the forty-four plates belonging to this subject, the first twenty-eight are 
coloured. 

[The Crustacea have not been studied. 

The Freshwater Malacostraca were undertaken by Prof. T. H. Huxley, 
but his contribution was never finished.] 

10. Araciinida Araneidea. Vol. I. : by the Rev. O. Pickard-Cambridge. 

This volume contains descriptions and figures of 417 new species of Araneidea, and 
a list of species identified by the author. The new forms are illustrated on the thirty- 
nine coloured plates. 

11. Arachnida Araneidea and Opiliones. Vol. II. : by F. O. Pickard-Cambridge. 

Vol. II. of this subject gives the systematic arrangement of the whole of the species 
of Araneidea described in Vol. I. and the large number added in Vol. II. A synopsis 
of the Families is given on pp. 541-544. The ' Opiliones,' or * Harvestmen,' are 
dealt with on pp. 546-585, and illustrated by three plates, LII.-LIV. The author is 
unable to say very much about the general distribution of the Araneidea, the material 



62 CONTENTS OF EACH VOLUME. 

at hand being insufficient for the purpose. They are said to date back at least as 
far as the Carboniferous period, ^vhen they were represented by forms of which the 
two species of LipMstius are the sole survivors. The earliest known Arachno- 
moiphid spiders in any way resembling those living belong to the Oligocene times. 
At that period there existed many forms, very similar to those occurring at the present 
time, whose remains have been found in amber washed up on the shores of the Baltic 
Sea. The Opiliones are stated to have preceded, during the Carboniferous epoch, the 
air-breathing scorpion, Anthrascorjnus, and their specialization probably began during 
the still earlier Silurian times, since in that period there existed an Arachnid which 
is a true scorpion in every sense, except that it had apparently no trace of air-breathing 
lung sacs. The ' Harvestmen ' make no web for the ensnaring of their prey, and the 
females, after depositing their eggs, take no further interest in their offspring. In the 
two volumes 1181 species are enumerated, this number including the 422 described 
as new in Vol. II. The Opiliones number 70 species, 58 of which are described as 
new, with 11 new genera. The ninety-three plates (39 in Vol. I. and 54 in Vol. II.) 
include figures of 981 species. 

12. Aeachnida Scoepiokes, Pedipalpi, and Solifug^ : by E. I. Pocock. 

Very little material was available for the study of these Arachnids, sixty-nine species 
only being enumerated for the three groups. The twelve uncoloured plates include 
figures of thirty-seven species. The Scorpiones are represented by three families, tiie 
Pedipalpi by two, and the Solifugse by a single family. 

In this Volume the author gives the geographical distribution under the heading 
for each genus, and no general ' Introduction ' to the whole subject was prepared, for 
want of data. 

13. Abachnida Acaeidea : by O. Stoll. 

'Jhe material for this subject was obtained almost entirely by the author during a 
residence of nearly five years in Guatemala. He made the drawings on the spot, but 
unfortunately he had no modern literature on Acarids with him, and his microscope 
was anything but satisfactory. Dr. Stoll's work, therefore, must be treated as a 
contribution to the fauna of Guatemala, rather than as an enumeration of the 
Acaridea inhabiting Central America. The types of the species described remained 
in his possession. In his ' Introduction' (published in 1893), p. vii, he states that it 
is remarkable that not one of the forms described represents a generic type entirely 
new or peculiar to the region. Doubtless a great deal remains to be done in the 
way of collecting before we shall have any true idea of the Acarid-fauna of Central 
America. 

The twenty-one coloured plates illustrate 43 species. 



ZOOLOGY. 63 

14. CniLOPODA and Diplopoda: by R. I. Pocock. 

The dates of publication of this Volume are, Chilopoda 1895-1896, Diplopoda 
1903-1910. For the two groups 255 species are enumerated, of which 106 are 
described as new, the Chilopoda numbering 53 (19 new) and the Diplopoda 202 
(87 new) respectively. The author, for want of sufficient data, does not give any 
particulars as to general distribution, beyond that mentioned under each genus or 
species in the text. The three plates belonging to the Chilopoda are partly coloured, 
the twelve others referring to the Diplopoda are uncoloured. 

[The Prototracheata [Peripatus, &c.) have not been studied. 
One species at least has been recorded from Nicaragua.] 

15. CoLEOPTERA. Vol. I. part 1 : by H. W. Bates : Cicindelidae and Carabidae. 

The author, who had previously studied the insect fauna of the Amazons during his 
long residence in that region, remarks, in his ' Introduction' (published in 1884), on 
the Central-American fauna of these two families as follows : — " The number of species 
(1086), belonging to 154 genera, is greater than the apparent poverty of tropical regions 
in Carabidae would have led us to expect. The tolerably well-worked valley of the 
Amazons, although rich in species of genera belonging to alluvial plains, and in 
arboreal forms, contains only 576 species belonging to 124 genera ; and the fauna of 
such tropical regions as the Malay archipelago is still poorer. The reason for the 
comparative paucity of Carabidae has been supposed, apparently on good grounds, to 
be that their place, as predaceous terrestrial insects, is to a great extent occupied by 
the ubiquitous ants. The undoubted fact that purely epigajous Carabidae, except 
marsh species, are scarce in the Tropics, especially near the Equator and in the low- 
lands, and that arboreal or climbing forms alone are numerous and varied, affords 
support to this hypothesis. The essentially Neotropical cliaracter of the Central- 
American fauna is generally admitted, and is strikingly confirmed by the Cicindelidae 
and Carabidae. But with regard to the northern limits of the fauna, and especially 
the extent to which Nearctic and North-temperate forms have penetrated the region 
from north to south, these are points not yet settled. Wallace included, or seemed 
inclined to include, the whole of the central highlands of Mexico and Guatemala in the 
Nearctic province, which must mean that the North-temperate American forms are 
there in the majority. The two families of Coleoptera we are dealing with do not 
support this conclusion. The Nearctic forms are comparatively few, and in the ' tierra 
templada' are far outnumbered by tropical genera. The northern limit of the Central 
American fauna appears to be — on the highlands, if not also on the maritime lowlands 
east and west — a little south of the political frontier of Mexico. Does the Central 
American fauna constitute one homogeneous province, or is it divisible into two 
subprovinces, as Salvin (• Ibis,' 1866, p. 202) has shown to be probably the case with 



64 COJSTENTS OF EACH VOLUME. 

the birds, many genera of which are represented by distinct species on each side of a 
line which he is inclined to place north of the Nicaraguan lakes and their outfall, the 
Rio San Juan 1 " 

'i'he thirteen coloured plates include figures of 324 species. 

Mr. Bates subsequently wrote two papers on the Mexican material received from our 
collectors afier the present Volume was closed : — 

(1) 'Additions to the Cicindelidae Fauna of Mexico' (Trans. Ent. Soc. Lond. 

1890, pp. 493-510, pi. 16). 

(2) ' Additions to the Cavabideous Fauna of Mexico ' (oj). cit. 1891, pp. 223- 

277, pis. 13, 14). 

16. CoLEOPTERA.. Vol. I. part 2: by I). Sharp; Haliplidse, Dytiscidas, Gjiiiiida', 

Hydrophilidae, Heteroceridae, Parnidse, Georissidoe, Cyathoceiida;, and 
Staphylinidae. 

For these nine families of Coleo])tera 1790 species are enumerated, 1405 of which 
belong to Staphylinidae, 168 to Dytiscidse, and 141 to Hydrophilida), the others being 
of limited extent. The author, in his 'Introduction' (published in 1887), states that 
the water beetles (Dytiscidas) are apparently subject to different laws of distribution 
from other Coleoptera, illustrating in this respect what he believes to be the case 
with aquatic organisms generally, viz. diminished endemicity, and therefore but little 
adapted for consideration in questions of zoo-geography. The study of the exotic 
Staphylinidae appears to be practically in its infancy, and but little can be done in the 
way of comparison with the fauna of other tropical regions. In the case of genera 
with a large number of species and a wide distribution, the Central American 
Staphylinids are stated to be much more nearly allied to those of South than to 
those of North America. Dr. Sharp is therefore inclined to agree with the opinion 
expressed by Bates that the fauna of the region under investigation is essentially 
Neotropical. The Staphylinidae first brought from the tropics included an undue 
proportion of comparatively large, bizarre, or brilliantly coloured forms, but it is clear 
that this was chiefly due to imperfect collecting. Indeed, one of the striking facts in 
connection with the material brought together by the Editors is the very large number 
of minute and obscure insects, so that it can scarcely be said that the Central American 
Staphylinidae are, on the average, larger or more biilliant than those of Europe. 

Of the nineteen coloured plates, fourteen are devoted to Staphylinidae, 443 species 
being figured altogether. 

17. Coleoptera. Vol. II. part 1: by Dr. Sharp, A. Matthews, and G. Lewis: 

Pselaphidae-Byrrhidae. 

The twenty-two families enumerated in this volume are collectively termed 
Clavicornia by various authors, and in recent catalogues of Coleoptera the Erotylidae, 



ZOOLOGY. 65 

Endomychidoe, and Coccinellidoe (these three families being here placed at the end of 
the Coleopterous series in Vol. VII.) are treated as belonging to the same division. 
The Silphidse, Corylophidse, Trichopterygidse, SphBeriids, and Scaphidiidae were 
worked out by Mr. Matthews, the Histeridae by Mr. G. Lewis, and the remaining 
sixteen families by Dr. Sharp, the last named author also contributing a list of the 
Khipidandri — a small group of somewhat uncertain position, but really belonging to 
the Tenebrionid-series near Boletophagini. The total number of species is 1629, of 
which 9'J6 are described as new. The dates of publication of the various subjects are : 
Pselaphidae and Scydmaenidae, 1887; Silphidae-Scaphidiidae, 1887-1888; Histeridse, 
1888; Phalacridse-Byrrhidae, 1888-1905; Khipidandri, 1905. The 'Introduction' 
to this Volume was simply an editorial note; but in his remarks on the Ilisteridse 
(p. 182) Mr. Lewis states that the chief interest of this series of insects centres 
in the species which feed on the wood-boring Coleoptera of other families. The 
nineteen uncoloured plates include, it is believed, some of the finest lithographic 
illustrations of beetles that have as yet been published, those devoted to the Colydiid* 
(plates xiv. and xv.) being particularly excellent. The artist, Baron Max Schlereth, 
was unfortunately unable to draw the whole number. 

18. Coleoptera. Vol. II. part 2 : by H. W. Bates : Pectinicornia and Lamellicornia. 

The total number of species for these two important families is 1100 — 72 Pectini- 
cornia and 1028 Lamellicornia. The author, in his ' Introduction ' (published in 
1890), remarks that the Pectinicorn-fauna is exceedingly poor in the chief family of 
the tribe, viz., the Lucanidse; but, on the other hand, it is unusually rich in the more 
aberrant family, the Passalidse. A comparison with such allied faunas from other 
parts of the world, which have been sufficiently worked out to give approximately 
accurate results, seems to show that the poverty in Lucanidae arises from Central 
America lying too far south to have been reached by many species of Old-World genera, 
and too far north for the genera characteristic of South Brazil, Chile, and the Andes. 
The conditions seem, however, to be very favourable to the Passalidse, which all pass 
their earlier stages in rotting tree-trunks, reaching their highest development and 
exhibiting more diversity of form here than in any other region. All the families 
of Lamellicornia are well represented in Central America ; they include 1028 species 
belonging to 127 genera, but a comparison of the Lamellicorn-fauna with that of 
other tropical regions of similar extent is impossible, as the necessary data do 
not exist in a connected form. The actual number of species described up to 1890, 
for the whole world, may be roughly estimated at 10,000. Of the twenty-four plates 
illustrating the 492 species figured, all but one (Pectinicornia) are coloured. 

Since this Volume was finished (in 1890) a good many species of Lamellicornia 
have been added by various authors, these showy beetles being great favourites 

BIOL. CE.NTR.-AMER., Introd. Vol., Jauuury 1915. K 



66 CONTENTS OF EACH VOLUME. 

with collectors, but the additions do not materially affect the conclusions arrived 
at by Bates. 

19. CoLEOPTERA. Vol. III. part 1: Serricornia: Buprestidae by C. O. Waterhouse ; 

Throscidae and Eucnemidte by G. H, Horn; Elaterid8e*-Dascillidje by 

G. C. Champion. 
For these families of the Serricorn-series, 1353 species are enumerated from Central 
America, 806 of which are described as new. The Buprestidae were published 
in 1882-1889; the Throscidae and Eucnemidae in 1890; the remaining families in 
1894-1897 ; and a short Appendix in 1897. The Buprestidae, numbering 434 species 
(exclusive of those mentioned in the Appendix), do not appear to be very much in 
evidence, apart from the gigantic Euchroma, in the tropical forests of Mexico and 
Central America, their place being to a great extent taken by the conspicuous Elaterid- 
genera Chalcolepidius and Semiotus. The open parts of Mexico, however, have a rich 
Buprestid-fauna, as shown by the number of species obtained by Hoge in his later 
expeditions to that country ; these were enumerated in the ' Supplement ' (published 
in 1889). The distribution of Conifers, which do not extend south of Nicaragua, 
probably affects the range of certain genera of this group. The Elateridae, numbering 
531 species, have exceedingly few endemic genera, and it may be said in a general 
way that the tropical forms are mostly confined to the forest regions, and that many 
of those inhabiting the open country or higher ground are nearly allied, or actually 
belong, to Nearctic genera. Pyrophorus has one species in North and sixteen in 
Central America, and is essentially Neotropical. The Dascillidae, with 130 species 
and twenty genera, is one of the ' neglected ' families of Coleoptera, and there are no 
available data for comparison with other regions. The Eucnemidae, of which a table 
of the genera is given by Horn (pp. 211-213), are represented by 113, the Throscidae 
by forty-four, the Cebrionidae by twenty-nine, and the Rhipidcceridae by fourteen 
species. 

The twenty-seven plates, one of which shows the form of the terminal segment of 
the males of the Buprestid-genus Fachyscelus, illustrate 648 species, 

20. Coleoptera. Vol. III. part 2 : by H. S. Gorham : Malacodermata. 

This Volume includes the following nine families : — Lycidae, Lampyridee, Tele- 
phoridae, Lymexylonidae, Melyridae, Cleridae, Ptinidse, Bostrychidae, and Cioidae. The 
total number of species enumerated is 813 ; but as very many of our specimens added in 
the ' Supplement ' were not critically examined by Mr. Gorham, it is probable that the 
actual number represented in the material obtained by our collectors is about 900. 
The author, in his 'Introduction' (published in 1886), states that it is now seen that 

• First nndertakeu by E. W. Janaon. 



ZOOLOGY. 67 

the tropical portions of the earth are as rich, or richer, in these groups (he presumably 
refers to Telephoridae) than the cooler parts. The Lycidse, Telephoridue, Lampyridae, 
and Cleridae clearly show a closer relationship between our fauna and that 
of Tropical South America than with any they possess with that of America 
north of Mexico — this being rendered even more apparent in such large genera as 
Calapteron amongst the Lycids and Chauliognathus in the Telephorids. More striking, 
however, is the marked peculiarity of the Central and South American genera as 
compared with the faunas of other parts of the globe. The author remarks : — " If it 
were true that similar conditions would produce similar forms, how is it that the 
specialized genera of the Neotropical regions are so dissimilar from those of the 
^Ethiopian and Indo-Malayan % Of the few genera common to Central America and 
to the Palsearctic or Tropical regions of the Old World, there is not one which is not 
of the rank of feebly differentiated forms, or persistent forms of an earlier stage of 
development. The total absence of many widely distributed well marked forms cannot 
be explained by any intrinsic dissimilarity in the conditions', which certainly do not 
differ more than they do in the wide areas over which such genera are dispersed, but 
rather by the isolation from these other areas, contact with which is only effected 
through the northern parts of the globe. One further fact is noted, viz., that, to 
whatever cause it may be due, where a genus is common to Central or South America 
and other distant parts of the World, it is also the case that it is represented by a 
species also identical or nearly related in the two districts. In this case its presence 
must be due to recent transmission, through artificial means, or to the conditions 
having long remained practically similar under which it exists in places so far apart." 

Thirteen coloured plates are issued in this Volume, 330 species being figured. 

Some of the Clerids received by us since the conclusion of Mr. Gorham's work have 
been described by Herr S. Schenkling in German publications, and a certain number 
of Bostrychidse have been dealt with or revised by M. P. Lesne. The Chauliognathinie, 
Malachiidse, and Melyridse, and a portion of the Ptinidse, have also been revised by 
Mr. G. C. Champion (Trans. Ent. Soc. Lond. 1913, 1914). 

21, 22. CoLEOPTERA. Vol. IV. parts 1 and 2 : by G. C. Champion : Heteromera. 

The fourteen families included in the Heteromerous-series are dealt with in two 
volumes : (1) Tenebrionidae, Cistelidae, Othniidse, Nilionidse, and Monommidae ; 
(2) Lagriidse, Melandryidse, Pythidae, QSdemeridae, Xylophilidae, Anthicidae, Mordel- 
lidse, RhipidophoridcB, and Meloidae. The general 'Introduction' (published in 1893), 
which includes a Table showing the geographical distribution of the genera repre- 
sented within the limits of Mexico and Central America, is given in part 1. The 
total number of species enumerated is 1776, of wiiich 1295 are treated as new. 
Amongst the Tenebrionidae, the apterous terrestrial forms clearly belong to the 

k2 



68 CONTENTS OF EACH VOLUME. 

Nearctic beetle fauna which extends down the central plateau to the Isthmus of 
Tehuantepec (Asida) or to the Los Altos region of Guatemala {Elceodes), while the 
winged forest forms are Neotropical. The winged genera inhabiting the open country 
(Ejntragus, Blapstinns, Xystropus, &c.) have a wide distribution, and such apterous 
forms as appear to be insensible to drought and heat (e. g. Zopherus) range from 
the Southern United States through our region to Colombia and Venezuela. The 
humid forest regions of Central America possess a very rich Tenebrionid-fauna, as 
so many species attack decaying trees, or the fungi growing upon them, even in 
gloomy places ; the Melandryidse, however, are poorly represented, as in other 
tropical regions. The Cistelidae, Xylophilidae, and OEdemeridae belong to the 
' neglected ' groups, not a single Central American species of any one of these families 
having been described before the publication of these volumes. The MelnidfB, on 
the contrary, had been studied by various Coleopterists, especially by E. Duges in 
Mexico, and many of the species were previously described. The Heteroraera, as 
stated in the Introduction, comprise a greater variety of forms than any of the other 
main divisions of the Coleoptera, nearly all of which are reproduced here. Probably 
no better case of mimicry or homochroism can be found amongst beetles than 
that existing between the Tenebrionid genus Ciiphotes and Cypher otylus (Erotylidse) 
and the Lagriid genus JJroplatopsis and Uroplata (Hispidse). Some of the smaller 
subcortical Cucujids of the Clavicorn-series are so closely related to various Pythids — 
from which they are only distinguishable by the number of joints in the hind tarsi of 
the males — that it is probable that these latter will have to be removed eventually 
from the heterogeneous series of families known collectively under the name 
lleteromera. 

The two volumes devoted to this group include forty-four coloured plates, 
illustrating upwards of 1000 species. 

23. Coleoptera. Vol. IV. part 3 : by D. Sharp and G, C. Champion: Curculionidae 
(part). 

The Rhynchophora, or weevils, include a larger number of species (3848) than any 
of the other main divisions of the Coleoptera, and no fewer than five volumes of the 
present series are devoted to their enumeration. In the First Volume (Part 3) 616 
species of the following subfamilies of the Curculionidae are dealt with : Attelabinae, 
Pterocolinae, Allocoryninie, Apioninae, Thecesterninae, and Otiorrhynchinse. Dr. Sharp's 
contribution, pp. 1-177, was published (pp. 169-177 excepted) in 1889-1891; this 
portion included the first five subfamilies and the apterous Otiorrhynchinse, and that 
of Mr. Champion, dealing with the winged Otiorrhynchinse, appeared in 1911. The 
Attelabinse and Apioninse, both numerous in species, do not differ greatly from the 
representatives of these subfamilies in temperate northern regions. The Pterocolinae, 



I 

ZOOLOGY. 69 

Allocoryninae, and Thecesteriiinae, each including a few species only, are common to 
North America and Mexico or Central America. The apterous Otiorrhynchids are 
mostly restricted to the central plateau — Eupagoderes, JEpiccerus, and IJparjriopsis 
heinij the dominant genera in the highlands of Mexico, — -while the winged forms 
])reponderate in the warmer forest regions to the east, west, and south, this 
distribution being similar to that of the Tenebrionids. Pandeleteius and Fantomorus 
are characteristic winged genera in the open country. Various papers on the 
Apioninae by Herr Hans Wagner, in which additional species from our region are 
described, have been published during recent years. Unfortunately, very little 
is known as yet of the actual food plants of any of the Central-American 
Rhynchophora. 

The fifteen coloured plates include figures of nearly 400 species. 

24. CoLEOPTEKA. Vol. IV. part 4 : by G. C. Champion : Curculionidse (part). 

The whole of this Volume is devoted to the Subfamily Curculiouinae, which are so 
numerous in the forest regions of Tropical America as to deter most Coleopterists 
from venturing to describe them. Twenty-two groups are enumerated, the Sitonina 
to the Cryptorrhynchina inclusive, numbering in all 1365 species, 1146 of which 
are treated as new. Some of the genera include a very large number of species : 
Conotrachelus (nearly 200), Anthonomus (over 100), Hilipus, Otuiocephalus, Cryptor- 
rhynchus, Enhidus, &c., so that it seems an almost hopeless task to prepare a complete 
list of these insects. Dr. Sharp and Mr. Champion have been the first to describe 
the whole of the Curculionidae of a tropical country, and, though the genera of the 
Cryptorrhynchina still remain in inextricable confusion, the present contribution will 
doubtless be of considerable assistance to future workers. Some of the Groups, 
Anchonina, Cholina, &c., are purely Neotropical ; others, Pissodina, Sitonina, Hyperina, 
Balaninina, Cleonina {Lixus excepted), belong to more temperate regions, but extend 
southward to within our limits. 

The thirty-five plates include figures of nearly 1000 species : nineteen (x. and 
xviii.-xxxv. inclusive) are coloured, one (xii.) partly coloured, and the rest 
uncoloured. 



25. CoLEOPTERA. Vol. IV. part 5 : by G. C. Champion : Curculionidae (continued). 

This Volume deals with four more Groups of the Subfamily flurculioninae — the 
Zygopina, Tachygonina, Ceuthorrhynchina, and Barina, the vast complex mass known 
as Barina being represented by eleven Sections, all but three of which are purely 
tropical. The Zygopina also are almost entirely tropical, a few forms only occurring 
north of the Mexican frontier and they are wholly wanting in the European fauna. 



70 CONTENTS OF EACP VOLUME. 

The Centhorrhynchina, on the other hand, are numerous in temperate regions and 
btit poorly represented in the tropics. Of the 908 species enumerated for the four 
frroups mentioned, 620 belong to Barina; 717 are described as new, with 81 new 
genera. 

The remaining groups of the Curculioninae — the Acamptina, Trypetina, and 
Cossonina— are described in Vol. IV. part 7, The twenty- three plates belonging to 
Part 5, illustrating 720 species, are wholly or partly coloured. 

26. CoLEOPTERA. Vol. IV. part 6 : Brenthidse' by D. Sharp ; Scolytidae by W. F. 

H. Blandford ; Anthribidae by K. Jordan. 

Three families of the Rhynchophora are enumerated in this Volume : the Brenthidie, 
published in 1895, by Dr. Sharp; the Scolytidae, published in 1895-1905, by 
Mr. Blandford ; and the AnthribidaB, published in 1906-1907, by Dr. Jordan. 
Part 6 was reserved for them in 1895, long before the Parts 4 and 5 were commenced. 
The general sequence of arrangement adopted in this work has therefore been 
interrupted by the interposition of these families in the present volume ; the 
remainder of the Curculioninae belonging to Part 5 were concluded in Part 7. The 
total number of species included in the three families is: Brenthidae 140 (104 new), 
Scolytidae 272 (181 new), and Anthribidae 193 (148 new). The Brenthidae are mostly 
confined to the forests of the tropics, and are particularly numerous in Tropical 
America, Madagascar, and the Indo-Malay region. As regards the Scolytidae the 
author, Mr. Blandford, remarks that the genera are either cosmopolitan or Neotropical, 
some of the latter being represented by stragglers in North America ; he also says that 
the southward limit of many northern forms is doubtless determined by the distribution 
of the Coniferae, which do not reach south of Nicaragua. Dr. Jordan, in his intro- 
ductory remarks on the Anthribidae (p. 299), writes as follows : — " The present 
treatise offers an illustration of the great increase in the number of known species 
when a tropical district has been more exhaustively examined. Hardly three dozen 
forms were known from Central America, while the material now before me comprises 
close on 200 species." These insects are rarely found in large numbers, owing to tlieir 
great resemblance to the bark of trees on which they occur. The author divides the 
Anthribidae into two subfamilies, the Pleurocerinae and the Anocerinse, and gives a 
key to the whole of the genera (pp. 300-302). In the text the type of each genus is 
indicated, and the species of the larger genera are tabulated. 

Fourteen plates are issued with this volume : three for the Brenthidae, six for 
the Scolytidae, and five for the Anthribidae, the last mentioned being coloured. 

27. CoLEOFTERA. Vol. IV. part 7 : by G. C. Champion : Curculionidaj (concluded). 
This Volume finishes the Rhynchophora, the rest of the Curculionidae belonging to 

the Subfamily Curculioninse — the Groups Acamptina, Trypetina, and Cossonina — and 



zooLOGr. .71 

the whole of the subfamily Calandrinse being here dealt with. An Appendix to the 
CurculionidsB is added, on pp. 178-212, enumerating a few additional forms and 
making some corrections to the synonymy. The Cossonina and Calandrinae are each 
represented in Central America by a large number of species, some of the Calandrids 
attacking palms, cacti, sugar-cane, maize, Musaceae, &c. The Mexican Cossonids (the 
genus Cossonus excepted) had been previously studied by Wollaston and the Calandrids 
by Chevrolat, nevertheless, with more abundant material, many new forms were found 
in our collection. Altogether .344 species are enumerated, 231 of which are described 
as new. The nine plates are coloured or partly coloured. 



28. CoLEOPTERA. Vol. V. : Longicornia by H. W. Bates ; Bruchides by D. Sharp. 

The greater part of this Volume (pp. 1-436), published in 1879-1885, is devoted to 
the enumeration of the Longicornia by Mr. Bates, the Bruchides, by Dr. Sharp, 
published in 1885, occupying pp. 437-504. Altogether the two tribes number 1423 
species: Longicornia 1273 (648 new), Bruchides 150 (117 new). Mr. Bates, 
in his Introduction to the Longicornia, published in 1886, remarks as follows : 
" Compared with the Tribe Geodephaga, it is beyond doubt far more numerously 
represented in tropical than in extra-tropical lands, and its species and genera are 
naturally multiplied in the highest degree in tropical forests, where woody vegetable 
growths, to which the Longicornia are almost exclusively attached in their larval states, 
are most numerous and varied. Although their beauty of form and colour has led to 
their having been industriously collected, it is evident, from the number of new 
species continually arriving from countries supposed to be fairly well explored, that we 
are as yet far from possessing even an approximately complete knowledge of the whole 
product of Nature in this department. This is partly due to the recondite and, to a 
great extent, nocturnal habits of a vast proportion of the species, and the difficulty of 
the search for them in dense primaeval forests where few clearings offer the necessary 
openings." The author thinks that the main conclusions arrived at after a similar 
examination of the Geodephaga are confirmed, viz. (1) that the Central-American 
fauna is essentially Neotropical ; (2) that the northern portion of the region (Mexico 
and Guatemala) is not an extension southward of the Nearctic Province, but (3) that 
it is a remarkably distinct subprovince of the Neotropical fauna. Dr. Sharp, 
in his remarks on the Bruchides, numbering 150 in all, says (p. 437-) that our 
knowledge of these insects is not sufficiently advanced to enable any trustworthy 
generalizations to be made in reference to the species found in Central America ; 
and, as the nortliern parts of Mexico had been inadequately explored, it was not 
possible to say what relationship existed between the North American species and 
those of the regions southward. 



<2 . CONTENTS OP EACH VOLUME. 

Of the twentj'-six coloured plates issued in the present volume, one only (pi. xxvi.) 
is devoted to the Bruchides. 

Two papers on the Longicornia, subsequently received from our collectors after 
Vol. V. was finished, have been published, entitled: "Additions to the Longicornia of 
Mexico and Central America": one by Mr. Bates (Trans. Ent. Soc. Lond. 1892, 
pp. 143-183, pis. v.-vii.) ; the other by Mr. Gahan (loc. cit. pp. 255-274, pi. xii.). 
These additions bring the total number of species of Central American Longicornia 
to 1372. 

29, 30. CoLEOPTERA. Vol. VI. part 1, and Supplement : by M. Jacoby : Phytophaga 
(pai-t). 

The enumeration of the Phytophagous Coleoptera was commenced in 1880, before 
the Salle collection had been acquired by the Editors and shortly after Mr, Champion 
had started on his journey to Central America. The collections made by him, and 
by Hoge in various parts of Mexico, increased our material so largely that a Supple- 
mentary Volume was added. Hence, two bulky Volumes, together including over 1000 
pages of text and forty-three plates, were required for the account of the Families 
Sagridse, Donaciidae, Crioceridae, Megalopodidae, Clythridse, Cryptocephalidae, Chlamy- 
didae, Lamprosomidae, Eumolpidae, Chrysomelidae, and Galerucidae. The Hispidae 
and Cassididae are described in Vol. VI. part 2. In the general Introduction to tlie 
two Volumes (Part 1 and the Supplement), published in 1892, the relative number 
of species for each family is shown in a Table, and the number of species of the 
larger genera {Diabrotica alone possessing 178, 116 of which are described as new, 
and Lema 129, with 73 new) is also noted. At the end of this Introduction, a 
systematic list of the whole of the species figured is given on pp. ix-xix. The Phyto- 
phaga are perhaps more in evidence, except in the denser forest districts, than any of 
the other families of Coleoptera in Tropical America, many of the species occurring 
in great abundance on the herbage in open places at the commencement of the rainy 
season. Some of them, like Orina in Europe, are extremely variable in colour, so that, 
as the author observes, nothing can be done by the systematic worker but to treat as 
distinct such forms as in his opinion differ sufficiently from their allies. Of the 2166 
species enumerated (in 1892) only about 90 are known from north of the Mexican 
boundary and about 150 from south of Panama : Central America is thus shown to 
have an exceedingly rich and peculiar Phytophagous fauna, of which the affinities are 
much greater with South than with North America. Since the publication of these two 
volumes, very little has been added to the Central American list beyoud a few forms 
described by Mr. Bowditch ; Mr. Gahan's papers on Diabrotica dealt with South 
American species only. 

About 1000 species are figured on the forty-three coloured plates. 



ZOOLOGY. 73 

31. CoLKorxERA. Vol. VI. part 2 : by J. S. Baly and G. C. Champion: Phytophaga 
(part). 
This Volume gives an account of the Hispidse by J. S. Baly, published in 1885-1886, 
with a short Appendix, in 1894, by G. C. Champion; and of the Cassidldae by 
G. C. Champion, published in 1893-1894. For the two families 453 species are 
enumerated — liispidae 226 (169 new), Cassididse 227 (55 new). The Central- 
American representatives of these groups are essentially Neotropical, America north 
of Mexico possessing a very limited number of species. Many of the characteristic 
South- American genera of Cassidida;, however, do not reach so far north as Panama, 
or, if present, are represented by very few species. Mexico is particularly rich in 
peculiar forms belonging to the genera Chelymorpha, Phijsonota, Coptocycla, &c. 
The Hispidae attack Musaceae, Bambusacese, &c., and abound throughout the warmer 
parrs of Tropical America. 

The thirteen coloured plates include figures of nearly 300 species. 

32. CoLEOPTERA. Vol. VII. : by PI. S. Gorham : Erotjlidae, Endomychidae, and 
Coccinellidae. 

The three families enumerated in this Volume are placed by recent writers in the 
Clavicorn-series, which are dealt with in Vol. II. part 1 of the present work. The 
total number of species is as follows: — Erotylida^ 282 (154 new); Endoraychidse 
81 (39 new); and Coccinellidae 239 (108 new). The Erotylidae, including the 
Languriides, were published in 1887-1889, the Endomychidae in 1889-1891, the 
Coccinellidae in 1891-1898, and a short Supplement to all three families, and the 
Introduction, in 1898-1899. The Author, speaking of the Erotylidae, exclusive of 
the Languriides, says that the members of this highly developed family of fungivorous 
beetles are largely endemic. The Endomychidae are somewhat poorly represented in 
Central America ; the Coccinellidae, or ' lady-birds,' on the other hand, are very 
abundant in species and individuals, the phytophagous Epilachnoe in particular. The 
small Aphid- and Coccid-devouring Coccinellids have of recent years become of 
economic importance. 

The thirteen coloured plates include figures of nearly 300 species. 



This is the last of the eighteen volumes devoted to the Coleoptera ; the total 
number of species enumerated is 18,029. Several genera of doubtful position, left 
undetermined by the various authors, have recently been described and figured, and 
the species of the genus Hapalips revised, by Mr. Champion (Trans. Ent. Soc. Lond. 
1913, pp. 58-169, pis. iii., iv.). 

33. Hymenoptera. Vol. 1. : By P. Cameron : Sessiliventria and Petioliventria. 

The fifteen families of Phytophagous or parasitic Hymenoptera worked out in this 
Volume number in all 1109 species, of which 596 are described as new. The 
BIOL, centr.-amer., Introd. Vol., January 1915. L 



74 'CONTENTS OF EACH TOrtTME. 

Sessiliventria incladft the Tenthredinidae, Ceplridae, Siricidae, and Oxyssidae, and the 
Petioliventria the CynipidiE, Figitidae, Chalcididae., Ichneumonidae, BraeoHidce, 
Stephanidae, Evaniidue, Trigonalidae, Peleoinidap, Proctotrapidte, and <Ohrysididae. 
When Mr. Cameron's work on these insects was published, in 1883-1888, scarcely any 
Mexican material was available for study, and the numerous IchneumonidiE described 
from that country in 1873 by Cresson could not therefore be identified. Mr. H. II. 
Smith subsequently made very extensive collections in Mexico of all the above- 
mentioned families, too late, unfortunately, to be included. Mr. Cameron's contri- 
bution, therefore, is very incomplete as regards the total num'ber of species inhabiting 
the region, althougli it adds considerably to our knowledge of these Hymenoptera. 
The numbers of species for the different families are as follows: Tenthredinidae 152, 
Cephidae 1, Siricidee 3, Oryssidae 2, Cynipidae 15, Figitidae 5, Chalcididae 121, 
Ichneumonidae 533, Braconidae 176, Stephanidae 5, Evaniidae 23, Trigonalidae 9., 
Pelecinidae 2, Proctotrupidae 43, and Chrysididae 19. 

The twenty coloured pilates include figures of 451 species. 

34. Hymenopteea. Vol. II. : by P. Cameren:: Fossores. 

In this Volume the species belonging to the Section Fossores »(the saT»fi- and wood- 
wasps), under which the author includes the FamHy Mutillidae, of the Hymenoptera 
Aculeata, are enumerated ; the other Sections are known as Heterogyna (ants), Diplo- 
ptery<:ia (Eumenidee and Vespidae), and Anthophila (bees). The Fossores, including 
Mutillidae, number 711 species, of which 391 are treated as new, the total number 
being: Sphegidae 67, Ampulicidae 2, Larridae 52, JSTyssonidas 32, Bembecidae 26, 
Philanthidae 54, Mimesidae 8, Pemphredonidae 3, CrabroniJae 24, Pompilidae 163, 
Scoliidae 59, Mutillidae 221. The pages 1-400 were published between 1888-1896, 
the rest in 1899 and 1900. Mr. H. H. Smith's Mexican collections .fortunately 
arrived in time to be included, so that the enumeration of the species in Vol. II. is 
more complete than in Vol. I. of the Hymenoptera. In the Introduction (published 
in 1900) there is a classified list of the 312 species figured on the fourteen xoloured 
plates. 

35. Hy>me\optkea. Vol. III..: by Prof. A. Forel : Heterogyna. 

Volume III. of the Hymenoptera, published in 1899-1900, contains an account, in 
French, of the Heterogyna or Ants, numbering 382 species, 66 of which are described 
as new. From the author's remarks on fpage 1. it would seem that a large number 
of the'Central-American forms are common to South America,.and that»the Formicid- 
fauna is mainly Neotropical. A few iclosely -related S. -American species are described 
in foot-notes. The four uncoloured .plates include figures of 57 species ; plate II. shows 
the nests of various -ants among leaves, or an the spines o{ Eri/t.hrina, Acacia, &c. A 
considerable number of species have been added to the fauna by Prof. Forel and other 
writers during the jpast ten years, showing that new forms still await the collector. 



ZOX)LOr»T. 75 

[The rest of the Aculeata, including the social-wasps, bees, etc., remain unwor&ecJ, 
ro contributor having been willing to undertake the large amount of material obtained 
in these groups. The late Lieutenant-Colonel Bingham commenced the study of the 
Diploptera shortly before his death, but nothing was published.] 

36-38. Lepidoptera Ritopamcera. Vols. I., II. (text),.III- (plates) : by F. D. Godmau 
and O. SalVin ; with (in Vol. II.) a Note on the Group EumsEidi by 
S. H. Scudder. 

Vol. I. of the Butterflies, published in 1879-1886, the' Introduction excepted, gives 
an account of the Nymphalida; — incltiding. the subfamilies Uanaina;, Satyrinse, 
Morphinse, Brassolint?, Acrseinoe, Heliconiina), and Nymphalinee, — Libytheidae, and 
Erycinidoe. Vol. II. includes the enumeration of the liycaenidae (published in 1887), 
Papilionidae, including the- subfamilies Pierinae and Papilioninae (1889-1893), and 
Hesperiidae (Hesperiinse, 1893-1899, Pamy)bilineB, by Godman olone, 1900-1901); a 
Suppkment to the whole subject by Godman, pp, 638-741 (L90.1)> and a Note on the 
Group Eumsjeidi, pp. 110-112, published in 1887, by Scudder, The Introduction to 
Vol. I. (pp. v-xlvi), by Godman, issued on the conclusion of Vol. II., in 1901, contains a 
description of the physical conformation of each of the countries belonging to tire Central- 
American region ; an explanation o.f the classification adopted ; remarks on the leading 
foi-ms belonging to each of the six families recognized (Nyinphalidae, Libytheida;, 
ErycinidcE, Lycaenid^e, Papilionidae, and Hesperiidse) ; a Table showing the geographical 
distribution of the genera (pp, xxii-xxvii) ;. particulaj-s as to the sources from whence 
the material was obtained, with ih-e names of the collectors and some remarks on the 
places visited by them ; and a systematic list of the whole of the species figured (1206) 
in the two Volumes (pp. xxxi-xlv), the plates numbering 112 in all. Altogether, 1805 
species (Nyniphalidte 588, Libytheid?e 1, Erycinidse- 240, Lyccenidse 234, Papilionidae 186, 
and Hesijeriidae 556) are enumerated, 376 of which are described as new, and 18 others 
from South America are added in foot-notes. In the Ilesperiidae the genitalia of most 
of the species are figured, many closely allied forms being definitely separable by these 
male-structures, find some of the species of the gtnus Fcqyilio are similarly illustrated. 

The study of these insects is said to prove conclusively (1) that the feuna is mainly 
a northern extension of that of Tropical South America, extending on the Pacific side 
to Mazatlan in Mexico and on the Atlantic side to a little beyond Ciudad Victoria 
in Tamaulipas, with many peculiarly modifi.ed forms in the region ; (2) that 
there are a considerable number of Nearctic genera and species coming down the 
central plateau a certain distance into Mexico, and some even to Guatemala ; (3) that 
there are no strictly alpine forms; (4) that the fauna of the Atlantic slope to perhaps 
as far south as Costa Rica is incomparably richer than the Pacific; (5) that some of 
the purely tropical genera do not reach north of Nicaragua, Costa Rica, or Panama. 
The publication of the concluding portion of the Rhopalocera, as in the case of the 

l2 



76 CONTENTS OF EACH VOLUME. 

Aves, was greatly delayed by the ill health and death of Salvin, in 1898 ; but he lived 
long enough to assist in completing the enumeration of the species of nearly all the 
families, with the exception of those of the subfamily Pamphilinae of the Hesperiidae, 
i. e. to p. 460 of Vol. II. 

Amongst the numerous papers on Neotropical Butterflies that have been published 
since the conclusion of Vol. II. (in 1901), one by Mr. H. H. Druce (P. Z. S. 1907, 
pp. 566-632, pis. xxxi.-xxxvi.) contains a revision of certain Central-American species 
of the genus Thecla. 

39-41. Lepidoptera Heterocera. Vols. I., II. (text). III. (plates) : by Herbert Druce : 
Sphingidse-Pyralidse. 

Vol. I. of this subject, published in 1881-1891, the Introductitm excepted, contains 
an account of the following 22 families of Moths : — Sphingidse, Castniidae, J^geriidai, 
Agaristidae, Zygaenidae, Arctiidae, Chalcosiidae, Lithosiidae, MelameridaE, Dioptidae, 
Liparidae, Saturniidse, Lasiocampidae, Limacodidoe, Bombycidae, Drepanulidae, Psychidae, 
Cossidae, Hepialidae, Notodontidae, Noctuidae, and Deltoidae. Vol. II., published in 
1891-1896, includes the Euschemidae, Uraniidae, Geometridae, Siculidae, and Pyralidae; 
and a very extensive Supplement, pp. 298-569, issued in 1896-1899, embraces all 
the families contained in both Volumes, another family, the Arbelidte (pp. 449, 450), 
being added. The author, in his Introduction to Vol. I., published in 1900, states 
that in 1880 less than 400 species of Heterocera had been recorded from Central 
America against the 3639 enumerated in these two Volumes. The Table given 
on page ix shows the number of species belonging to each of the 28 Families and 
of those peculiar to the region. He says that it is almost impossible to give any 
generalizations as to the composition of the Central American Heterocera, because so 
little is known of the species inhabiting the adjacent parts of South America ; but it 
may safely be said that their affinities are almost entirely South American, and the 
few northern forms that do occur mostly belong to widely distributed genera. 

The 101 coloured plates include figures of 1926 species, a complete list of which is 
given in the Introduction to Vol. I., pp. xi-xxxi. 

Since the conclusion of Mr. Druce's work, in 1900, a large number of species have 
been added, especially from Mexico and Costa Rica, mainly by Mr. W. Schaus, who 
has specially visited Costa Rica, &c., in search of Lepidoptera. Sir George Hampson, 
too, in his revision of the Noctuidae, Pyralidae, &c., has also contributed largely to our 
knowledge of the Tropical American Heterocera, the critical study of which had 
scarcely been commenced when Vol. I. was undertaken. 

42. Lepidoptera Heterocera. Vol. IV. (text and plates); by Lord Walsingham : 
Tineina, Pterophorina, and Orneodina. 

This Volume, commenced in 1909, contains the enumeration of the Tineina, 



ZOOLOGY. 77 

Pterophorina, and Orneodina, and a few supplementary genera and species omitted in 
the other volumes dealing with the Heterocera. The Tineina are grouped under 22 
Families — Lavernidae, Gelechiadae, CEcophoridse, Ethmiadae, Biastobasidse, Stenomidie, 
[.Egeriadfe], [Cossidae], Arrhenophanidae, Sparganothidae, Olethreutidee, Tortricidae, 
Phaloniadae, Carposiiiidae, Hemerophilidae, Coleophoridae, Heliodinidae, Hypono- 
meutidiB, Fhyllorycteridae, Tineidae, Acrolophidas, and Nemophoridae. Two of them, 
the ^geriadae and the Cossidie, were previously dealt with in detail by Mr. Druce 
in Vol. I., their true affinities with the Tineina not having been recognized at 
that time. Lord Walsingham, in his Introduction, alludes to the great assistance 
he has received from Mr. J. H. Durrant in the preparation of this Volume, which 
is illustrated by ten coloured plates, representing 350 species of Tineina. 

43. DiPTER.\. Vol. I. : by Baron C. R. Osten Sacken, the Supplement by S. W. Wil- 

liston, J. M. Aldrich, W. M. Wheeler, and A. L. Melander : Cecido- 

myiidse-Empidse. 
Twenty-four Families of Diptera are enumerated in this Volume — Cecidomyiidae, 
Mycetophilidae, Bibionidae, Simuliidse, Blepharoceridae, Culicidae, Chironomidae, 
Psychodidae, Tipulidae, Rhyphidae, Stratiomyidae, Tabanidae, Chiromyzidse, Leptidae, 
Xylophagidae, Acanthomeridae, Mydaidae, Nemestrinidae, Bombyliidae, Therevidae, 
Cyrtidae, Asilidue, Dolichopodidae, and Empidae. Osten Sacken's work was published 
in 1886-1887. The Supplement by the American writers Williston, Aldrich, Wheeler, 
and Melander was issued in 1900-1901, and the editorial Introduction in 1901, on 
the conclusion of the whole subject. The material examined by Osten Sacken was 
very meagre, and though more specimens from Mexico were available for the 
Supplement, yet it was insufficient to give more than a general idea of the Dipterous 
fauna of Central America. The present Volume, therefore, is merely a contribution 
towards the study of these insects, rather than a complete list of the forms actually 
inhabiting the region. Altogether, 977 species are enumerated, 330 of which are 
described as new. The six coloured plates illustrate 120 species. 

During recent years the Mosquito fauna of Panama has been specially studied in 
the canal-zone by Mr. A. Busck, and his report has been published [Smithsonian 
Contributions, vol. lii. pp. 49-77 (May 1st, 1908)]. Many new species of Culicidae 
from Panama and other places in Central America have been described by Theobald, 
Dyar, and Knab. 

44. Diptera. Vol. II.: by F. M. van der Wulp: CEstridae, Muscidae, and Hippo- 

boscidse. 

This Volume contains the enumeration of the numerous groups of Diptera known 
collectively under the name of Muscidae, for which abundant Mexican material collected 
by Mr. H. H. Smith was avaihible, but the work was only about half finished when the 



78 CONTEMS or EACH VOLUME. 

author died in 1899. No other contrihutor could be found to undertake the groups 
Agromyzirae, Borborinsp, Chloropsirse, Drosophilinse, Ephydrinae, Geomyzina?, and 
Sapromyzinae, of the Muscidaj Acalypterse, and the Phoridaj, and the volume was 
closed as it stood. Pages 429-489, containing the Hippoboscidic and the Supple- 
ment to the other families, Were publ shed after v. der Wulp's death, although 
the new species of Muscidae Calyptrae had already been described by him in the 
'Tijdschrift voor Entomologie' for 1 892. Altogether, 1095 species were named by the 
author, of which 585 were treated as new, the publication of his work extending from- 
1888-1900, but the Titlepage and Introduction were not issued till 1903. 

The thirteen coloured plates include figures of 287 specie*. Some of the Trypetinse 
are figured on the plaiJes, others (11) in the text. 

45. DiFTERA. Vol. III. : by S. W. WiUiston : Syrphklse, Conopids^, PipuHculidae, and 

Platypezida;. 

Prof. Willi ston's contribution to this Volume, pp. 1-^89, was published in 1891- 
1892, and the list of subsequently described species, and the general ind«x to the- 
whole of the three volumes, pp. 93-127, in 1903. The material again was scanty in 
comparison with the large number of species which must exist in the region, but 
among the Syrphids the genera Baccha, Volucella, and Eristalis were particularly 
well represented. The total number of species enumerated is 325-, of which (17 
are new. The two coloured pfates includte figures' of 29 species. 

46, RHTJfCHOTA Heteroptera. Vol. I. : by W. L. Distant : Pentatoniidae', Coreidae, 

I^ygseidae, Pyrrhocoridae, and Capsidoe. 

Mr. Distant's enumeration of the above mentioned families of Rhynchota Hetero- 
ptera (pp. 1-303) was issued in 1880-1884,. the Supplement tO'the same (pp. 304-351) 
in 1889-1893, and the Appendix (pp. 452-462) in 1893, the delay in publishing 
the Supplement being duo to his long absence in South Africa. Altogether, 
1108 species are recognized, more than half of which are described as new. In the 
Introduction, published in 1893, the number of genera belonging to each family is 
given and the geographical distribution shown in short Tables. Amongst the 
Xrygseidse nearly half the genera are stated to be endemic. The Pentatomidce include 
377 species and are, therefore, very well represented in Central America, though 
many of the genera are common to the Neotropical region. The Capsidoe, too, with 
313 species, are very numerous, but the tropical representatives of this family are 
but little known as yet, and no comparison with the fauna of the adjacent portion 
of South America can be made. The thirty-nine coloured plates include figures of 
vipwards of 900 species, a complete list of them being given in the Introduction, 
pp. xi-xx. 



•ZOOLOGY. ;'f9 

47. RiiYNCiiOT.^. Heteroptera. Vel. II. : by G. C. Champion : Tingitidae-Corixidse. 

Mr. Champion here deals with nineteen families of Rhynchota Heteroptera (making 
twenty-four in a;ll), twelve belonging to the Gymnocerata and seven to the Crypto^ 
.cerata. The number of species enumerated is 592, nearly half of which are 
•described as new. Fifteen additions to Vol. I., described or recorded by various 
authors in other works, between 1893 and 1901, are noted on p. S84, the total 
number of Meteroptera enumerated in the two Volumes being 1715. The Tingitidoj, 
Aradidae, Reduviidae, and Anthocoridae are abundantly represented in Central America, 
rthe respective numbers for each being 78 (66 new), 78 (40 new), 204 (78 jaew^), 
and 54 (32 new). As with the five families belonging to Vol. I., no comparison is 
(made with the fauna of the adjacent regions southward for want of sufficient data. 
JSIost of the genera belonging to the aquatic Cryptocerata are, in common with, other 
■water-insects, more or less widely distributed. The twenty-two plates, six of whicli 
.are coloured, include figures of 513 species, a complete list of which is given in the 
■Introduction, pp. xi-xvi. The Index for the two volumes is included iu Vol. JI„ 
pp. 3«5-416. 

A8. KaYNGHQTA 'HoMOPTER.\. Vol. I. : by W. L. Distant and W. J^. Fowler: Cicadadae- 
Delphacida". 

Mr. Distant's first contribution to this Volume, on the Cicadidae and Fulgoridaq, 
•pp. 1-41, was published in 188,1-1887, the Supplement to the same, pp 42, 43, in 
a.900, and the Appendix, pp. 140-146, in 1905, his vi'ork having been interrupted 
during long absences from England. The rest of the volume, dealing with three 
additional species of Dictyopharinae, the Flatidse, Derbidae, Cixiid*, Achilidoe, Issidaej 
:and Delphacidae, by the Eev. W. W. Fowler, pp. 44-139, 146, 1^7, appeared ia 
1900-1905. Apart /from.the Gicadidte and Fulgoridae, -very little appears to be known 
.about the South American members of the above mentioned families, and nearly all the 
Central American.representatiyes are described as new by Mr. Fowler. Altogether 323 
species are enumerated, 208 of which are treated as new, with twenty-five new genera. 
The thirteen coloured ,plates include figures of .267 species, ja coniplete .list oi" which 
is given in the Introduction, pp. vii-ix. Four species of Cicadidae are also figured 
in the text, -on pp. 140-143. A few cCeotral American .Cicadidae have since beeu 
described by Mr. Distant, in other publication?. 

49. Rhvnchota Homoptbea. Vol. H. >Part 1, by W. W. ^Fowler : Membracidce- 
G.yponidae. Part 2, by T. D. A. Cockerell : Aleurodidae and CoccidiE. 
Part 1 of this Volume contains the enumeration, of foixr more .families of Homopterfi 
— the Membracid«e, Cercopidse, Tettigoniidae, and Gyponidae, 659 species in .all 
(exclusive of the 29 noticed in the Supplement, some of which are doubtful), 385 of 
of these being described as .new, with 42 new genera. .Pages 1-316 were published 



80 CONTENTS OF EACH VOLUME. 

in 1894-1903, and 317-322 in 1909, the conclusion of the Volume having been 
delayed for several years by the American contributor who had volunteered to 
determine the Jiissidae, Bythoscopidae, &c. The MSS. and material not being forth- 
coming, the volume was closed as it stood. The twenty-one coloured plates, ten 
of which are devoted to the extraordinary insects known as Membracidae, include 
figures of 494 species, a list of which is given in the Introduction, pp. vii-xi. The 
Index for Vols. I. and II. is also issued in this volume, pp. 323-339. The Supplement, 
pp. 318-322, contains notes on the 29 species added by Buckton and others during 
the progress of the work. 

Part 2 of this Volume, by Prof. T. D. A. Cockerell, issued in 1899 (the separate 
Index, pp. 35-37, was not published till 1909, when Volume II. was closed), gives 
a list of the 161 Aleurodidae and Coccidse from Mexico and Central America known 
to the author at that date. Eleven species and two genera are described as new, 
and nine forms are figured in the text. With these insects, the total number of 
Homoptera recorded from Central America is 1143. 

[The few specimens of Thysanoptera obtained have been examined, and some of 
the species described, by Mr. R. S. Bagnall (Journ. Linn. Soc, Zool. xxx. pp. 369- 
387, i)ls. ii., iii., 1910). Mr. D. L. ('rawford has also recently written various papers 
on ' Thysanoptera of Mexico and the South' (Pomona Joutn. Eut. i., ii. 1909-1910).] 

50. Neuroptera : Ephemeridse by A. E. Eaton ; Odonata by P. P. Calvert. 

The Rev. A. E. Eaton's account of the Ephemeridae, based upon very few specimens, 
was published in 1892 ; that of the Odonata by Prof. P. P. Calvert, with abundant 
material, in 1901-1908 *. Altogether 308 species are enumerated : Ephemeridse 32 
and Odonata 276. Prof. Calvert's Introduction, pp. v-xxx (published in 1908), 
mainly consists of two Tables: (A) " Alphabetical List, by countries, of the localities 
in which the Odonata were collected," and (B) " Systematic list and distribution of 
the species and varieties," 293 in all ; and a coloured map to show the actual distri- 
bution of mean annual temperatures. Table A gives the name of the Department, 
Territory, or State to which each locality belongs, the elevation of these places in 
metres and feet, the names of the collectors, the dates of capture, general remarks, &c. 
Table B shows the geographical distribution and the temperature zones in which 
the species occurs in each country, this being further illustrated by the map. The 
composition and ecological relations of the Odonate fanua of Mexico and Central 
America have been discussed by Mr. Calvert elsewhere [Proc. Acad. Nat. Sci. Phil. 
(1908) 1909, pp. 460-491, pi. xxvi.]. He has since visited Costa Rica, May 1st, 
1909-May 9th, 1910, to collect data on (1) the seasonal distribution of the Odonata, 

* The Odonata were undertaken successively by Mr. McLachlau and Prof. Karsch, before Prof. Calvert 
proceeded with the work. 



ZOOLOGY. 81 

(2) their larval forms, and (3) their habits. An account of his journey was published 
in the 'Entomological News' for July 1910, pp. 334-337, and this has been followed 
by other papers on the results of his researches, the most important being the 
discovery of Odonate larvaj found living in the water which collects between the bases 
of the leaves of Bromeliads growing upon trees, and of the rearing of the imagines of 
Mecistogaster modestus from these larvae. 

The ten plates belonging to this Volume, of which three are coloured, include 
figures of 178 species, all but nine appertaining to the Odonata. 

[No contributor has been found for the remaining families belonging to the 
Neuropterous-series in the wide sense, viz. Trichoptera, Neuroptera Planipennia, 
Mallophaga, Psocidge, Termitidae, Embiidfe, and Perlidse.] 

51. Orthofteea. Vol. I. : Forficulidae by Count de Bormans ; Blattidae, Mantidae, 

Gryllidae, and Locustidae by Dr. Henri de Saussure, assisted by Dr. Leo 
Zehntner and A. Pictet. 

Five families of Orthoptera are dealt with in this Volume : the Forficulidae by 
de Bormans (published in 1893), and the Blattidae, Mantidae, Gryllidae, and Locustidae 
byde Saussure (published in 1893-1899). The Introduction, including the systematic 
list of species figured, was issued in 1900, when Vol. II. of this subject was com- 
menced. De Saussure, in his enumeration, also considered it necessary to describe, 
notice, or figure various allied South-American or Antillean forms. Under the 
heading to each family, tribe, &c., he deals with the general classification of the 
Neotropical fauna, and gives synopses of the genera, and under the genera them 
selves dichotomous keys to the species. The total number of species recognized as 
belonging to Central America is 633, of which 224 are new : Forficulidae 41 
(8 new), Blattida; 156 (60 new), Mantidje 61 (33 new), Gryllidae 102 (43 new), 
Locustidae 273 (80 new). The twenty-two plates, of which, iii., vi., vii,, and viii., 
are coloured, include figures of 310 species. 

52. Orthoptera. Vol. II. : Acridiidae by Prof. Lawrence Bruner (the subfamily 

Tettiginae by A. P. Morse) ; Phasmidse by R. Shelford. 

This Volume includes the enumeration of the two remaining families of tbe 
Orthoptera. The Phasmidae should have been placed between the Blattidae and 
Mantidae, in Vol. I., but the account of them had to be deferred until the appearance 
of Brunner v. Wattenwyl's and J. Redtenbacher's Monograph, the concluding portion 
of which was not published till 1908. Prof. L. Bruner's work on the Acridiidae, in 
which Mr. A. P. Morse assisted him with the subfamily Tettiginae, was issued in 
1900-1908, the Tettiginae having appeared in 1900-1901. The list of the Pliasmidie, 
published in 1908, was compiled from the above mentioned Monograph, the authors 

BIOL. CENTR.-iVMER., Introd. Vol., June 1915. M 



82 CONTENTS OF EACH VOLUME. 

of which included the material obtained by the Editors of the present work. As in 
Vol. I., the species, excepting the Tettigina?, likely to occur in Mexico or Central 
America are included in the enumeration. Altogether 663 species are regarded 
as belonging to the fauna, although in many cases, owing to lack of material, there 
is at present no actual record from within the limits of Central America. The 663 
species are apportioned thus: Acridiidae, including the 26 Tettiginae, 522 (138 new) ; 
PhasmidiE 141. The Index to the two volumes occupies pp. 379-412 in Vol. II. 
The eight uiicoloured plates (four for each family) include figures of 120 species, some 
of which are Nearctic or Neotropical forms closely related to those known to inhabit 
C'entral America. 

BOTANY. 

53. Vol. I. : by W. B. Hemsley, with a Commentary on the Introduction and Appendix 
by Sir J. D. Hooker. 

This Volume contains the enumeration of the Polypetala;, Ranunculacese- 
Araliaceae, which was issued in 1879-1881; the Preface, Introduction, and 'Com- 
mentary ' appeared in October 1888, on the completion of the entire subject. When 
this took place, the temporary titlepage and the Index to Vol. I. (pp. 577-619), 
issued in October 1881, were both replaced in October 1888, the former by a titlepage 
to include notice of the ' Commentary ' by Sir J. D. Hooker, and the latter 
by a general index for the whole of the four Volumes of text, this appearing in 
Vol. IV. pp. 333-498, 1887-1888. In the Introduction, pp. ix-lxi (issued in 1888), 
Mr. Hemsley deals with geographical distribution under the following headings : 
(1) Statistics of the Phanerogamic Flora of the world ; (2) Statistical comparison of 
the Floras of large and widely separated areas ; (3) Generic and specific composition 
of the Floras of diff'erent areas ; (4) The distribution of some of the largest Natural 
Orders ; (5) The Primary Botanical regions of the World considered in their relations 
to the Zoological regions ; (6) Comparison of the Zoological with the Botanical 
regions; (7) Outlying Australian types of vegetation; (8) Botanical division of the 
earth into primary regions. Sir J. U. Hooker in his ' Commentary on Mr. Hemsley's 
Introduction and Appendix to the Botanical part of the Biologia Central!- Americana' 
(pp. Ixii-xlviii) also gives his own views on the Botanical kingdoms, under these 
headings: (i.) The North Temperate Kingdom of the Old World; (ii.) The Tropical 
Kingdoms of the Old and New World ; (iii.) The three Southern temperate regions 
(Extratropical x\merica, Africa, and Australia). 

54. Vol. II. : by W. B. Hemsley. 

The Gamopetalae, Caprifoliacese-Plantaginese, are dealt with in this Volume. Tlie 
temporary titlepage and the Index to Vol. II., pp. 577-621, both issued in June 



BOTANY. 83 

1 882, were cancelled in 1888, the " Contents" being added to the title and the Index 
replaced (as in Vol. I.) by one for the complete series of four volumes, this being 
inserted at the end of Vol. IV. 

55. Vol. III. : by W. B. Hemsley ; the Cycadaceae by W. T. Thistelton-Dyer. 

The Incorapletae, Monocotyledones, and Cryptogamic Vasculares, Nyctaginse- 
Rhizocarpge, are enumerated in Vol. III. The Cycadaceae, pp. 190-195, were studied 
by Mr., now Sir W. T. Thistelton-Dyer, and his portion of the subject was issued 
in 1883. 

56. Vol. IV. : by W. B. Hemsley. 

•This Volume, the last of the series, exclusive of that occupied by the Plates, 
includes : (1) The Supplement to Vols. I.-III. ; (2) The enumeration of a small 
collection of plants made in Cozumel Island in 1885; (3) A list of plants from 
Holbox, Mugeres, Cozumel, and Ruatan Islands made in 1886 ; (4) Additions to the 
list of Costa Rican ferns, bringing the number up to 134 ; (5) Appendix. The 
Appendix deals with a variety of subjects : (i.) Preliminary remarks ; (ii.) A sketch of 
the history of the botanical exploration of Mexico and Central America ; (iii.) Outlines 
of the geography and the prominent features of the Flora of Mexico and Central 
America ; (iv.) Summary and analysis of the Flora ; (v.) Relationships with the Floras 
of other regions ; (vi.) Further details of the distribution of some of the more 
prominent natural Orders ; (vii.) A specimen of the mountain Flora of South Mexico 
and Central America ; (viii.) Altitudinal distribution of Orchids in South Mexico 
and the dominating features of the general vegetation; (ix.) Recapitulation of the 
dominant features of the Flora of Mexico and Central America, and remarks on its 
probable derivation ; (x.) Bibliography. The general index, as stated above, occupies 
pp. 333-498. The total number of species enumerated in the four volumes is 
11,626, of which 196 are indicated as new by the authors. 

57. Vol. V. : by W. B. Hemsley. 

The 111 Plates (including XLI.*), eighteen of which are coloured, together form 
Vol. V. of this subject. A complete list of the 144 species figured is given on 
pp. v-viii. Plate CX. consists of a Map of Mexico and Central America, showing the 
then known northern limits (in 1887) of the Phanerogamic epiphytes, the Coco-palm, 
the Marcgraviacese, the Vochysiaceae, and the southern limit of Pines. 



M '2 



84 CONTENTS or EACH VOLUME. 

ARCHEOLOGY. 

58, 59. Vol. I. (text and plates) : by A. P. Maudslay. 

The Volume of text contains the Preface and Introduction to the entire subject, 
and a general account of the ruins at Copan. The latter are described under five 
headings: (1) Principal Notices and descriptions of the Ruins; (2) Personal narrative; 
(3) General description of site ; (4) Detailed description of principal structures ; 
(5) Description of Stelae and Altars. In addition to the 119 folio plates, there are 
.numerous illustrations in the text. The text of Vols. I.-IV. is arranged for binding 
in one Volume; the plates illustrating the subject form four separate volumes, 
when bound. 

58, 60. Vol. II. (text and plates) : by A. P. Maudslay. 

The Ruins described in Vol. IT. are those of Quirigua, Ixkun, Yaxche, Rabinal, 
Chacujal, Utatlan and Iximche, Mixco (foundation-mounds), and Menche. These 
places are all shovpn on the 98 folio plates, the first of which is a map of Guatemala 
and the adjoining countries, showing the position of the ruins ; and there are also 
various illustrations in the text. 

58, Gl. Vol. III. (text and plates) : by A. P. Maudslay. 

The Ruins described in Vol. III. are those of Chichen Itza and Tikal. The 82 folio 
plates, the first of which is a map of Yucatan and the country to the south of it, show 
these places, and there are also three illustrations in the text. 

58, 62. Vol. IV. (text and plates) : by A. P. Maudslay. 

The Ruins of Palenque are described in Vol. IV. under eight headings : (1) Personal 
Narrative ; (2) Principal Notices and Descriptions of the Ruins ; (3) General de- 
scription of the site; (4) Detailed description of the principal structures; (5) Pottery; 
. (6) Initial Series or date ; (7) The two-headed dragon ; (8) The Water-plant. These 
subjects are shown on the 93 folio plates. 

63. Appendix (text only) : The Archaic Maya Inscriptions : by J. T. Goodman. 

Pages 1-149 of this Volume are occupied with Mr. Goodman's account of these 
inscriptions; and pp. 151-264 by the Annual, Chronological, and Perpetual Calendars 
(Tables), and the Working Chart. The ' Appendix ' forms the entire Part VIII. of 
the Archaeology, and is unaccompanied by Plates. 



[ 85 ] 



SUMMARY OF CONTENTS OF EACH VOLUME. 





Total 

number of 

species. 


New 

species 

described. 


New 

genera 

described. 


Plates 

(including 

maps). 


Species 
figured. 


Pages of 
letterpress. 


Dates 

of 
issue. 


1. 


Introductory Volume 








10 




viii & 149 


1915 




Zoology. 
















2. 
3. 
4. 
5. 
6. 
7. 


Mammalia 


181 
il413 

695 


14 
58 


1 
3 


22 

84 
76 


34 

149 
138 


XX & 220 

xliv & 512 

iv & 598 

iv & 510 

vii 
XX & 326 


1879-1882 
1879-1904 
1888-1904 
1897-1904 
1879-1904 
1885-1902 


Aves. Vol. I 


„ A'^ol. II 


Vol. Ill 


Vol. IV 


Eeptilia and Batrachia 


8. 
9. 


Pisces 


415 

871 


16 
81 


1 
1 


28 
44 


78 
333 


xxiiii & 203 
xxviii & 706 


1906-1908 
1890-1901 


Terrestrial and Fluviatile Mollusea 


10. 


Arachnida Araneidea 


1181* 


f417 
t422t 


54 


39 


}98l{ 


XV & 317 


1889-1902 


11. 


„ „ and Opiliones. 


38 


54 . 


X&610 


1897-1905 


13. 
14. 


Arachnida Scorpiones, Pedipalpi, 
and Solifugse 


69 


8 


. . 


12 


37 


i&71 


1902 


Arachnida Acaridea 


55 
255 


40 
106 


12 


21 
15 


43 
139 


xxi & 55 
i&217 


1886-1893 
1895-1910 


Chilopoda and Diplopoda 


15. 


Coleoptera. Vol. I. part 1 .... 


1086 


384 


8 


13 


324 


X&316 


1881-1884 


16. 


„ Vol. I. part 2 


1790 


1470 


103 


19 


443 


xvi & 824 


1882-1887 


17. 


Vol. II. part 1 


1629 


996 


83 


19 


475 


xii & 7r7 


1887-1905 


18. 


Vol. II. part 2 


1100 


494 


19 


24 


492 


xii & 432 


1886-18f)0 


19. 


Vol. III. part 1 . . . . 


1353 


805 


28 


27 


648 


xvi & 690 


1882-1897 


20. 


Vol. III. part 2 .... 


813 


535 


28 


13 


330 


xii & 372 


1880-1886 


21. 
22. 


Vol. IV. part 1 

Vol. IV. part 2 .... 


|l776 


1295 


92 


f 23 
21 


568 
500 


xxxiv & 572 
X&494 


1884-1893 
1889-1893 


23. 


Vol. IV. part 3 


016 


516 


45 


15 


397 


vi & 354 


1889-1911 


24. 


Vol. IV. part 4 


1365 


1146 


69 


35 


997 


viii & 750 


1902-1900 


25. 


Vol. IV. part 5 . . . . 


90S 


717 


81 


23 


720 


viii & 514 


1906-1909 


26. 


Vol. IV. part 6 . . . . 


605 


433 


32 


14 


324 


iv & 396 


1895-1907 


27. 


Vol. IV. part 1 .... 


344 


231 


17 


9 


275 


vi & 221 


1909-1910 


28. 
29. 
30. 


Vol. Y 


1423 i 
.2166 


765 
1363 


57 
43 


26 
43 


515 
1010 1 


xii & 526 

XX & 625 

iv & 374 


1879-1880 

1880-1892 
1888-1892 


Vol. VI. part 1 .... 
Vol. VI. part 1, 

Supplement . . 


31. 


Vol. VI. part 2.... 


453 


224 


4 


13 


291 


X&250 


1885-1894 


32. 

33. 


Vol. VII 


602 
1109 


301 
596 


21 
21 


13 

20 


287 
451 


xii & 276 
xii & 488 


1887-1899 
1883-1900 


Hymenoptera. Vol. I 


34. 


Vol. II 


711 


391 


1 


14 


312 


xi & 413 


1888-1900 


35. 


Vol. Ill 


382 


66 




4 


57 


ii & 170 


1899-1900 


36. 
37. 


Vol. II. 


|l805 


376 


119 






xlvi & 487 
iv & 782 


1879-1901 
1887-1901 


38. 


Vol. III. 








112 


1206 


iv 


1879-1901 


39. 

40. 


„ Heterocera. Vol. I. 
Vol. II. 


13639 


1312 


41 




.. • 


xxxii & 490 
iv & 622 


1881-1900 
1891-1900 


41. 


Vol. III. 




, , 




101 


1920 


iv 


1881-1900 


42. 


Vol. IV. 


973 § 


586 


54 


10 


350 


xii & 482 


1909-1915 



• Araneidea 1111, Opiliones 70. t Araneidea 364, Opiliones 58. 

+ Longicornia 1273 (648 new), Bruchides 150 (117 new). 

§ Exclusive of 52 species previously enumerated by Druce in Vols. I. and II. 



86 



SUMMAEY OF CONTENTS OF EACH VOLUME. 



Total 

number of 

species. 



43. 
44. 
4o. 

46. 
47. 

48. 

r 

49.< 

I 

L 



SC- 
SI. 
52. 



53. 

54. 
55. 
5H. 
57. 



58.' 

59. 

60. 
61. 
62. 
63. 



Zoology (cont.). 

Diptera. Vol. I 

Vol. II 

Vol. Ill 

Ehynchota Heteroptera. Vol. I. 

Vol. II. 
Ehynchota Homoptera. Vol. I. . . 
Vol. II. 

part 1 

Vol. II. 

part 2 

Neuroptera : Ephemeridas .... 

„ : Odonata 

Orthoptera. Vol. I 

Vol. II 



Total 



BOTAMr. 



Vol. 



I. 

II. 
III. 
IV. 

V. 



Aech^olout. 



Vol. I. (text) 

„ II. (text) 

„ III. (text) 

„ IV. (text) 

Vol. I. (plates) 

„ II. (plates) 

„ III. (plates) 

„ IV. (plates) 

Appendix (text onlv). 



977 
1095 

320 
1108 

607 

323 

659 

161 
32 
276 
633 
663 



38637 



New 
species 
described. 



330 
585 
67 
581 
289 
208 

385 

11 

8 

77 

224 

138 



19067 



( 

11,626^' 

I 



22 

135 

34 

5 



New 

genera 

described. 



11 
40 
1 
105 
30 
25 

42 



1 
14 
26 



1373 



Plates 

(including 

maps). 



6 
13 

2 
39 
22 
13 

21 



1 
10 

22 
8 



1173 



111 



119 
98 
82 
93 



Species 
figured. 



120 

287 
29 
903 
513 
267 

494 



91 
109 I 
310* 
120* 



18051 



Pages of 
Ittterpress. 



viii & 378 
X &490 
viii & 127 
XX & 462 
xvi Si, 416 
X&147 

xii & 339 

ii&37 

XXX & 420 I 

X&458 
viii & 412 



144 



Dates 

of 
issue. 



Ixviii & 576 J 

iv <fc 576 t 

W& 711 

iv & 498 

viii 



viii & 69 
iv & 47 
iv&60 
iv & 38 

iv 

iv 

iv 

iv 
xii & 264 § 



1886-1901 
1888-1903 
1891-1903 
1880-1893 
1897-1901 
1881-1905 

1894-1909 

1899 

1892 
1901-1908 
1893-1899 1 
1900-1909 

1879-1915 



1879-1888 

1881-1888 
1882-1888 
1886-1888 
1879-1888 



1888-1902 
1899-1902 
1895-1902 
1896-1902 
1889-1902 
1899-1902 
1895-1902 
1896-1899 
1897 



* Including allied South- or North-American or Antillean forms figured for comparison. 

t The Introduction, including the systematic list of species figured, was not issued till April 1900, when 
Vol. II. was commenced. 

J The Index (pp. 577-619 of Vol. I. and pp. 577-621 of Vol. II.) was replaced, on the completion of 
Vol. IV., by a fresh one for all four volumes, which is placed at the end of IV. 

§ Calendars, pp. 151-264, not paged. 



Six maps, in addition to the eight included in the Introductory Volume, have been issued in various volumes 
of the present work : two, showing the " Distribution of Freshwater Fishes in Mexico and Central America " 
(Pisces, Introduction, 1908) ; one, illustrating the Distribution of pines, coco-palm, etc. (Botany, Vol. V., 
plate 110) ; one, giving the " Actual Distribution of Mean Annual Temperatures," illustrating the distribution 
of the Odonata (Neuroptera, Introduction, 1908); one of Guatemala and the adjoining countries, showing 
the position of the ruins in Mexico, Guatemala, aud Honduras, described by Mr. Maudslay (Archfeology, 
Vol. II., plate 1) ; and one of Yucatan and the country to the south of it, showing the position of other 
places described by the same author (Archteology, Vol. III. plate 1). 



[ S7 1 



ORIGIN, ETC., OF THE FAUNA AND FLORA OF 
CENTRAL AMERICA. 



[The following summary and conclusions on the origin and geographical distribution 
of the Mammalia, Reptilia, Batrachia, Pisces, Arachnida, Chilopoda, Uiplopoda, and 
Prototracheata of Central America, and the faunistic divisions of the region, have been 
kindly supplied by Messrs. R. I. Pocock and C. Tate Regan ; and Mr. W. B. Hemsley 
has given us a similar account of the Flora. These valuable contributions are based 
upon our present knowledge of the subject, bringing the account up to date. As 
regards the Insecta, which occupy so large a portion of the work, no satisfactory con- 
clusions can be formed till they have been more thoroughly collected and studied in 
other tropical regions, and perhaps till we know more of them in a fossil state. — Ed.] 

FAUNA. 

MAMMALIA. 

By R. I. Pocock, F.R.S. 

Since the publication of Mr. E. R. Alston's volume on the Mammalia of Central 
America in 1882, great advances have been made in our knowledge of the subject. 
The past history of many of the orders and families has been more or less accurately 
ascertained by palseontological research, principally in the United States and in the 
Argentine ; and the modern methods of collecting and preserving existing material 
have led to the discovery of large numbers of genera, species, and subspecies, and 
in addition have thrown such light upon the vertical and horizontal distribution of 
American Mammals, as a whole, that it has become possible to map their zones and 
provinces with much greater precision. 

This progress in accuracy of information has been accompanied by a gradual change in 
the conception of systematic terms, with the result that what our predecessors ignored 
as " local varieties " are now regarded as " species " and, by a logical sequence, the old- 
time species are being given the rank of genera. Quite apart, too, from the names 
that have been introduced to designate new forms, considerable changes have taken 
place in the nomenclature of long-established species and genera. Opinions may differ 
as to the advisability and advantage of the alterations coming under these headings, but 
they have to be admitted and dealt with by faunistic and systematic workers. 



88 OKIGIN, ETC., OF THE FAUNA. 

In view of the additions and modifications thus briefly referred to, it is necessary to 
gather together the principal records that have been published during the last quarter 
of a century concerning the Mammals of Central America *. The plan adopted will 
be to treat the orders separately and to append to each a brief account of its palseon- 
tological history and such particulars of its distribution as have a bearing on the 
matter in hand. 

Order PRIMATES. 

The American Monkeys (Cebidse) and Marmosets (Callitrichidae) constitute the 
Platyrhine tribe of Primates as opposed to the Catarhine tribe embracing the 
Monkeys, Apes, and Men of the Old World. The Central American species belong 
to genera inhabiting also South America, where they are restricted to the forested 
districts east of the Andes, most of them being of tolerably wide range within those 
limits. In Central America they are similarly limited to the forest. Of the Cebidse, 
Squirrel Monkeys {Saimiri=Chrysothrix) extend to Costa Rica, Spider-Monkeys 
{Ateles), and Howlers (Alouatta) to Vera Cruz in Mexico, Capuchins (Cebus) to 
Nicaragua, and Douroucoulis [Aotus—NyctipUhecus) to Costa Rica and Nicaragua, 
but the latter locality needs confirmation. Of the Callitrichidse, Geoffroy's Tamarin 
{(Edipomidas geoffroyi) ranges from Colombia to Costa Rica. 

There are no Monkeys in the Antilles, apart from introduced species, and none in 
North America. 

There is a good deal of evidence that Monkeys are the descendants of Lemurs, which 
date back to the Lower and Middle Eocene of North America and Europe, and survive 
at the present time in Africa, Madagascar, and Southern Asia. Oddly enough, this 
group of Primates never seems to have entered South or Central America, and no 
fossil remains of Monkeys have been found in North America. Nevertheless, Platy- 
rhine Monkeys go back to the Upper Miocene in South America if the reference of 
Homunculus to that group be, as it appears to be, correct, and no extinct members 
of the group (which is, as a whole, more Lemuroid than the Old World Catarhini) 
have been found outside that continent. There are one or two facts which suggest 
Africa as their original home. 

In Madagascar there has been found an extinct Lemur (ArchcBolemur) which is 
claimed to have Platyrhine characters in its jaws and teeth, and the Oligocene of Egypt 
has yielded a Monkey (Parapithecus) structurally bridging the interval between existing 
Lemuroids of the Tarsioid (Tarsius) group and the Simiid or monkey-like Primates. If 
the interpretations put upon these fossils be correct, it seems that the transitional stages 
between the Lemurs and the Monkeys were probably evolved in an Afro-Mascarene 

* It is perhaps necossary to explain in this connection that here and elsewhere in this article the term 
"Central America" is used comprehensivelj-, as it is used in the title of the 'Biologia,' to embrace the area lyiug 
between, and including, Mexico and Panama, Lower California being excluded. 



MAMMALIA. 89 

continent. Coupling this inference with the absence of extinct Monkeys in North 
America and of Lemurs in South America, the view that the ancestors of the Platy- 
rhini entered South America from Africa is at all events provisionally defensible. 

Order INSECTIVORA. 

America is not rich in members of this order. Two families, the Talpidae (Moles) 
and Soricidae (Shrews), occur in North America. Although the Talpidae range as far 
south as the Southern States, they do not appear to enter Mexico or Central America. 
The Soricidae, on the contrary, are represented in Central America by the three genera 
Sorex, Cryptotis (sometimes regarded as a subgenus of the North American Blarina), 
and Xotiosorex. Of these, Sorex has a wide range over Europe, Asia, and North 
America, and passes at high altitudes through Mexico to Guatemala; Notiosorex 
inhabits the Southern States of North America (Texas etc.) and Mexico (Sinaloa, 
Jalisco). Cryptotis, which replaces in Central America the North American genus 
Blarina *, ranges mostly at high altitudes through Central America (Mexico, Yucatan, 
Guatemala, Nicaragua, Costa Rica) to Colombia, Venezuela, and Guiana. 

Shrews related to Sorex and Blarina have been recorded from the Oligocene of 
Europe and North America, and Sorex itself goes back to deposits of that age in Europe, 
appearing in North America in the Pleistocene, and since no Insectivores of this group 
occur in South American deposits f , it seems clear that the Central American species 
are southern migrants from North America. 

Order CHIROPTERA. 

The Central American genera of this order, fifty or more in number, are assigned 
to eight families — the Emballonuridae, Noctilionidae, Phyllostomida;, Desmodontidas, 
Natalidae, Thyropteridae, Vespertilionidaj, and Molossidae $. Geographically these 
families may be referred to two categories : the first containing the Vespertilionidae, 
MolossidfE, and Emballonuridae, which are represented in the Old World as well as in 
America ; and the second containing the remaining five, which are mostly restricted to 
Central and South America, a few genera only occurring in the Southern States of 
North America. Of this second category, by far the most important numerically are 
the Phyllostomidae, the American Leaf-nosed Bats §, the genera of which, occurring in 

* A species of Blarina has been recorded from Costa llica, but there seems to be some doubt about the 
correctness of the localit)'. 

t The only extinct South American Insoctivore known is Necrolestes of Upper Miocene age, the nearest 
ally of which appears to be the Golden Mole (Chrysochlorls) of South Africa. 

X Pending the completion of Dr. Knud Andersen's ' Monograph of the Chiroptera,' in course of publication 
by the Trustees of the British Museum, the classification and nomenclature here followed are those of Miller 
(Bull. U.S. Nat. Mus. Ivii. 1907). 

§ The Leaf-nosed Bats of the families Ehinolophidce, Hipposiderida;, and Megadermidae are restricted to 
the Old World. 

BIOL. CENTK.-AMER., Introd. Vol., June 1915. N 



90 OEIGIN, ETC., OF THE FAUNA. 

Central America, are about equal numerically to the genera of all the other families 
put together. 

A few only of the more important genera need be mentioned. 

The Vespcrtilionidse are cosmopolitan to the limit of tree-growth. Three familiar 
Old World forms occurring in our area are : Myotis, extending from North America to 
the Argentine ; Eptesicus, spreading as far south in America as Guatemala ; and Fip}- 
strellus, reaching South Mexico. Corynorhinus, the American representative of the 
Old World Plecotus, ranges from North America into Mexico. Other American 
genera are : Lasypterus, extending from North to South America ; Rhogocessa, from 
Central Mexico to Venezuela; and Bceodon, known only from Jalisco, in Mexico. 
Finally, Antrozous, formerly, but wrongly, affiliated with Plecotus and ranging from the 
Southern States of the Union to Central Mexico, constitutes with the Australasian 
genus Nyctophilvs a special subfamily of the Vespertilionidse. 

The Molossidae are more southern in distribution than the Vespertilionidoe. In the 
Old World their northern limit is South Europe and Asia. One genus only, Nyctinomus, 
found also in Europe, Africa, Asia, and Australia, reaches America, where it spreads 
from the Southern United States, though Central America, to Chili. Of exclusively 
American genera, Promops and Molossus pass from Mexico into South America, while 
Eumops has a northern range into the Southern States. 

Of the Emballouurida;, mainly a tropical family, not one of the Old World genera 
occurs in America, and the American genera (e. g. Saccopteryx, Peropteryx, and 
Diclidunis) extend from Central into South America. 

Of the families confined to America, the Noctilionidae contains two genera — one South 
American, the other (Noctilio) passing northwards to South Mexico ; the Natalidse and 
Thyropteridae contain one genus each, namely Natalus and Thyroptera, ranging 
respectively from South America to Central Mexico and Honduras ; the Phyllostomidte, 
divisible into several subfamilies, have, broadly speaking, a distribution similar to that 
of the other exclusively American families just mentioned, though some of the genera 
have a wider, others a narrower, range. For example, of the Chilonycterinse, Morinoops 
reaches Texas, Chilonycteris and Pteronotus do not go north of Mexico ; of the 
Phyllostominae Xenoctenus is only known from Costa Rica, Glyphonycteris passes 
southwards from that country to Peru, whereas Otopterus extends from the Southern 
States of North America to Guatemala ; of the Glossophaginae, Choeronycteris reaches 
from South America to Arizona. The single species of Ilalonycteris is known only from 
Costa Rica, while Lichonycteris, also monotypical, has been recorded from Nicaragua 
and Guiana. Similarly, in the Stenoderminae, the monotypical Centurio and Ectophylla 
are Central American, but the other Central American genera range into tropical 
South America. Finally, the family Desmodontidse, or true Vampyres, differing from 
the Phyllostomidae by their large shear-like anterior teeth, is represented by two 
penera, Desmodus and Liphyllus, which are distributed from Southern Mexico to 
Paraguay and Brazil. 



MAMMALIA. 91 

The small number of extinct Chiroptera known, i. e. a few genera belonging to the 
Hipposideridae and Vespertilionidae from Eocene and Oligocene deposits of Europe, 
throw no light upon the present distribution of the order. 



Order RODENTIA. 

In the number of species and genera inhabiting Central America, this order rivals, 
if it does not surpass, the Chiroptera, the four usually admitted suborders (Sciuro- 
morpha, Myomorpha, Hystricomorpha, and Lagomorpha *) being well represented. 

Suborder Sciuromoepiia. 

Two families of this suborder have to be considered, namely the Petauristidse or 
flying Squirrels and. the Sciuridae or true Squirrels, Ground-Squirrels, and Marmots. 
The one American genus of Petauristidae, Sciuropterus, ranges from Europe and 
Asia into North America, where it extends from Alaska to Guatemala. Of 
the Sciuridae, the Chipping Squirrel {Eutamias) and the Souslik (Citelius) have a 
distribution as extensive as that of Sciuropterus, whereas an allied form, Callospermo- 
philus, and the Prairie Marmot {Cynomys) are restricted to the South and Western 
States of North America and North Mexico. True arboreal Squirrels are represented 
by vast numbers of species and subspecies in North, Central, and South America, as 
well as in Europe, Asia, and Africa ; but the genera and subgenera to which they have 
been referred require revision and re-definition before their mutual affinities can be 
accurately ascertained. Of the Central American genera, Sciurus (s. s.) appears to 
be the only one that occurs in the Old World, where it ranges from Japan to Ireland. 
In America it is generally distributed throughout the States, occurs everywhere in 
Central America at high and low levels, and enters the northern countries of South 
America (Colombia, Ecuador). An allied form with many species, Guerlinguetus 
(Parasciurus), is credited with a range from the United States to Peru. Others that 
may be mentioned are Baiosciurus (extending from Mexico to Nicaragua), Syntheo- 
sciurus (known only from Chiriqui in Panama), and Microsciurua (alleged to spread 
from Costa Rica to Peru f ). 

Genera regarded as primitive Sciuromorphs were abundant in North America 
(Ischyromyid;r) and Europe (Pseudosciuridae) during early Tertiary times (Eocene and 
Oligocene), extinct Marmots [Palmarctomys) and the still existing Cynomys have been 
traced back to the Upper Miocene in North America, while Sciurus itself has survived 
in Europe and North America since the Upper Oligocene. Smce no extinct forms are 

* Apart from the Lagomorpha, these suborders are not perhaps susceptible of precise definition. They 
are here maintained for the sake of convenience. 

t Beavers (Castoridaj) should perhaps bo added to this section, since they range in North America at least 
as far south as Sonora. In the Old World, Beavers occur in Northern Asia and North aud Central Europe. 

n2 



92 OEIGIN, ETC., OF THE EAUNA. 

known in South America, unless of quite recent date, it seems clear that Central 
America received this prominent element of its fauna from North America. 

Beavers also seem to have originated in North America, where they date back to 
the Middle Oligocene, but Castor itself appeared in Europe in the Pliocene and in 
North America in the Pleistocene. 

Suborder Myomokpha. 

The Central American Myomorphs belong to the three families Muridsc, Hetero- 
myidse, and Geomyidse, the two last-named being restricted to America. 

The genera of Muridae belong to the three subfamilies Microtinae, Neotominse, and 
Cricetinae. Of the MicrotinEe, Microtus (s. s.) extends over the greater part of Europe, 
Asia, and North America, and enters Mexico (Puebla, Jalisco, Oaxaca). Two sub- 
genera are known only from Central America, namely, Ortlmomys from Oaxaca and 
Herpetomys from Guatemala, both at high altitudes. The other genus Pitymys has a 
singular distribution, ranging in the Old World from South and Central Europe to 
Asia Minor (the Mediterranean Area), and through the Eastern States of North 
America to Mexico (Vera Cruz). 

Of the Neotominse, Xenomys, Hodomys, Nehonia, and Teanopus are confined to 
Mexico, while Neotoma, represented by a host of species, overlaps thera to the north 
and south with a range from the Southern States of North America to Guatemala. 

The Cricetinae have an extraordinarily wide distribution, namely Africa, including 
Madagascar, Europe, Asia, and practically the whole of America. The Central 
American genera are too numerous to mention in detail. Some occur in North and 
Central America, others in Central America alone, others are common to Central and 
South America, while others extend over the three areas mentioned. To the last 
category belong Eeithrodontomys, Oryzomys, and Siymodon, ranging from the Southern 
States of the Union to Ecuador, Chili, and Peru, respectively. As examples of the 
first group may be cited Peromyscus [Ilesperomys) distributed from Labrador to 
Panama, and Baiomys and Onychomys from the Southern States to Guatemala and 
Panama. Peculiar to Central America are Nyctomys and Ototylomys, ranging from 
Mexico to Panama and Guatemala, and Siijmodontomys known only from Costa Rica, 
while Akodon, extending all over South America from Costa Rica, and Nectomys, 
occurring in Nicaragua, Ecuador, and Colombia, are representatives of genera common 
to Central and South America. 

The genera of Heteromyidae belong to two subfamilies, the Heteromyinse and 
Dipodomyinae. Of the latter, Perodipus and Lipodomys (Jerboa-Mice) are restricted 
to the South and South-western States of North America and to Mexico (Zacatecas, 
Vera Cruz, etc.). Of the former, Ileteromys, Perognathus, Liomys, and Chcetodipus 
occur in the Southern United States and Mexico, the first passing southwards into 



MAMMALIA. 93 

Colombia and Venezuela. The Geomyidse, or Pocket-Gophers, are mainly Central 
American, Thomomys, Cratogeomys, and Geomys ranging into the Southern States of 
the Union, Platygeomys, Zygogeomys, etc., being Mexican, Orthogeomys extending 
from Mexico to Guatemala, and Macrogeomys occurring from Nicaragua to Panama. 

The palsBontological history of the Voles (Microtinse) before the Pleistocene is some- 
what meagre, but Microtiis has been traced back to the Pliocene in Europe and to the 
Pleistocene in North America. Eecords of the Gricetinse are more complete, and 
extend back to the Lower, Middle, and Upper Oligocene in Europe and North America, 
the existing Feromyscus dating in North America to the Miocene. In South America 
Cricetines appear to be of recent (Pleistocene) date. The Heteromyidae and Geo- 
myidsB, closely related families, seem to have originated in North America, where 
they have been traced respectively to the Lower and Upper Oligocene. 

The palaeontological evidence, therefore, suggests that the ancestors of the Central 
and South American Myomorpha came down from the north. This is no doubt true 
of the Geomyidae and Heteromyidoe, but it is not so certain of all the Cricetines. The 
failure to discover fossil remains of small rodents such as these in mid-Tertiary 
(Upper Miocene) deposits in South America cannot be regarded as conclusive evidence 
of their absence there at the time the deposits were laid down; moreover, the 
occurrence of genera of this subfamily in Madagascar strongly suggests their existence 
in Africa at a sufficiently early date to permit their migration thence into South 
America by the land-connection which there are other grounds for thinking may have 
joined these two continents together. 

Suborder H Y s T R i c o M o R p H A. 

Three families — the Octodontidae, Dasyproctidae, and Hystricidae — occur in Central 
America, but neither is peculiar to the country, though the Dasyproctidae are found 
elsewhere only in South America and the West Indies, None of the Central 
American genera of these families enters the United States. 

Of the Dasyproctidae, Dasyprocta (Agouti) ranges from Brazil through Panama to 
Mexico, probably Vera Cruz ; Agouti or Coslogenys (Sooty Paca) also passes from 
Panama to Vera Cruz, The Hystricidse are represented by Coendou (:= Synetheres), 
the common South American Tree-Porcupine, which extends from Bolivia and Brazil 
to the mountains of Mexico. In North America its place is taken by Erethizon, 
distributed from Alaska and Canada to Arizona. One other genus of this family, 
ChcBtomys, occurs in South America only. These arboreal Porcupines constitute the 
subfamily Erethizontinae as opposed to the Hystricinae or Ground- Porcupines of the 
Old World. The Octodontidae, generally distributed througliout South America, 
comprises a large number of genera, two of which, Loncheres and Proechimys, enter 
Central America, the latter as far as Nicaragua, the former only to Panama. A third, 
Hoplomys, occurs in both these countries. 



94 OEIGIN, ETC., OF THE FAUNA. 

Although South America must be regarded as the present headquarters of the 
Hystricomorphs, on account of the number and diversity of the genera that occur there, 
the group is not restricted to America. There are several African genera belonging 
to or closely allied to the Octodontidse ; and Porcupines {Ilystrix, Atherura) are found 
in Tropical Asia as well as Africa. 

The palseontological history of the Hystricomorphs is very different from that of 
the other suborders of Eodentia. Genera regarded as ancestral members of the group, 
and constituting the family Theridomyidse, were living in Europe from the Middle 
Eocene to the Upper Oligocene, and true Porcupines, related to the existing species of 
Hystrijc, were in Europe in the Middle Miocene ; but none of these extended into 
North America. On the other hand, genera assignable to the Octodontidae and 
Erethizontine Porcupines suddenly appear in South America in Upper Miocene 
deposits — that is to say, considerably later than the date of the first known appearance 
of the group in Europe. The available evidence therefore points to the conclusion 
that this suborder made its way into South America from Africa, and subsequently 
spread northward into Central America and into North America, EretMzon dating 
from the Pleistocene in the latter country. 

Suborder LAGOMORPnA. 

The Ochotonidse (Picas) — which occur in Central Asia and spread through Western 
North America from Alaska to California, Utah, and Montana — do not enter Mexico. 
The Leporidse have a much wider range. Three genera have been recorded from 
Central America, namely Lepus, Sylvilagus, and Homerolagus. The last is mono- 
typical, and has been discovered at Puebla, Popocatepetl, etc., in Mexico at a height 
of 10,000 to 12,000 feet. Sylvilagus runs right through Central America from the 
Southern United States into South America. Lepus, in the modern sense, on the other 
hand, does not go south of Mexico (Durango, Tamaulipas, Potosi, etc.), whence it 
spreads northwards to Greenland and is also found in Asia, Europe, and Africa. 

The earliest palaeontological records of Hares (Leporidae) are from the Lower 
Oligocene of North America {Palceolagus). Lepus itself was in existence in the Upper 
Oligocene of that continent, and appeared in Europe in the Lower Pliocene. It seems 
therefore that this suborder of Rodents was evolved in North America, and thence 
made its way into the Old World and into Central and South America. 

Order CARNIVORA. 

The terrestrial or fissiped Carnivores of Central America belong to the five families 
Canidse, Procyonidse, Ursidse, Mustelidae, and Felidae. 

Canidce : — The genus Canis is represented in North America by many species or 
local races belonging to groups typified by C. occidentalis and C. latrans, corresponding 



MA.MMALIA. 95 

respectively to the Wolves and Jackals of the Old World. Both groups occur on 
the Mexican plateau — distinct species of the latrans or Prairie Wolf type having 
been recorded from Tamaulipas, Durango, Puebla, etc., as far as Guatemala, and an 
unknown form is said to occur in Costa R.ica. A second genus, Urocyon, commonly 
regarded as a Fox, extends from the Eastern States of North America, through 
Central America (Mexico, Yucatan, Guatemala), to Colombia in South America. 

There are reasons for thinking that Urocyon is more nearly allied to the South 
American group of Canidse typified by Cerdocyon thous than to Vulpes. If this be so, 
the genus constitutes an interesting geographical link between the Canidae of North 
America and those of South America. In the latter continent there is a very large 
number of species ranging from the extreme north to Cape Horn and the Falkland 
Islands. These are referable to several groups possibly of generic status ; but none 
of these can be definitely affiliated with Canis or Vulpes or other genera inhabiting 
North America and the Old World. The apparent absence from the tropical countries 
and highlands of Central America of Canidse related to the forest and Andean species 
of South America is a singular fact. 

Mustelidoe. — The dominant Central American Mustelids appear to be Skunks, which 
are represented by Mephitis ranging from Canada to Guatemala, Spilogale from the 
Southern United States through Mexico to Costa Rica, and Conepatus with much the 
same northern limits as Spilogale, but extending throughout South America to Tierra 
del Fuego, mostly at high levels. Weasels {Mustela) range all over Europe, Central 
and Northern Asia, North America, and thence southwards through Central America 
into the northern countries of South America. The remaining terrestrial forms, th« 
Tayra {Tayra =■ Galera) and the Grison { G risen =: Galictis), are more southern in 
distribution. The former is found in Vera Cruz, Nicaragua, Panama, Guiana, Peru, 
Brazil, etc., and the latter in Honduras, Yucatan, Costa Rica, and thence southwards to 
Patagonia. Finally, Otters (Lutra) are represented in Cental America by two species, 
one described from Jalisco, the other from Nicaragua. This genus is practically 
cosmopolitan in the Old World, apart from Madagascar and the Australian Region, 
its exceptionally wide range being possibly connected with independence of 
terrestrial barriers resulting from its power of making its way by swimming along 
coast-lines. 

Felidoe. — Cats are abundant everywhere, and belong to many distinct groups, probably 
of generic value. The short-tailed or Lynx-group, which is widely distributed over 
Northern Europe, Asia, and North America, enters Mexico (Tamaulipas, Sinaloa). The 
Puma-group, which has no close kinship with Lions and Leopards of the Old World, 
ranges from the confines of Canada to Patagonia and occurs in Central America at both 
high and low levels from Mexico to Panama. The other species of this family — 
belonging to a variety of groups akin to the Jaguar {F. onca). Ocelot {F. pardalis), 
Eyra (F. eyra) groups, etc. — are related to the South American species of those 



96 OEIGIN, ETC., OF THE EAUNA. 

names. Both Ocelots and Jaguars frequent the forested districts of South America, 
and pass through Central America into the United States north of Mexico. There is 
no doubt that the Jaguar is closely allied to the Leopards, and is the only American 
member of the genus which comprises also the Lion, Tiger, Leopard, and Ounce 
of the Old World. Ocelots, too, are not apparently generically separable from 
several of the Eastern Asiatic species, and there appears to be a form closely akin to 
them in tropical West Africa. 

ProcyonidcB. — The five admitted American genera of this family occur in Central 
America, which in this respect is richer than either the northern or southern moieties 
of the New World. Procyon (Raccoon) has a range almost equal to that of the Puma 
in its extension from near the borders of Canada to Paraguay. Subspecies akin to 
the typical North American form {lotor) reach Southern Mexico ; two species appear 
to be restricted respectively to Yucatan and Panama, while the South American 
P. cancrivorus enters the latter State. A peculiar species of Kasua (Coaiti Mondi) 
occurs on Cozumel Island, Yucatan, the rest of the Central American forms recorded 
from Mexico (Colima, Chihuahua), Yucatan, Guatemala, and Panama being regarded as 
subspecies of the South American N. narica; Bassariscus (Cacomistl) ranges from 
Texas and Oregon through Mexico to Guatemala and Panama ; liassaricyon is 
represented by species from Nicaragua, Costa Rica, and Panama, and occurs as far 
south as Ecuador ; while Potos, better known as Cercoleptes (Kinkajou), ranges 
from Mexico (Vera Cruz) Costa Rica, and Guatemala into Colombia, Ecuador, 
Venezuela, etc. 

In the Old World the family is represented by JElurus (Panda) of the Eastern 
Himalayas and probably by ^luropus of Moupin in Eastern Tibet. The latter, 
however, is claimed by some authorities to be Ursine in its affinities. It shares the 
characters of the Ursidae and Procyonidae, and may perhaps be regarded as the living 
link between the two fiimilies. 

XJrsidce. — Bears akin to the North American Black Bear {TIrsus {Euarctos) ameri- 
canns) extend into Mexico as far as Coahuila and Chihuahua. South of Mexico the 
family is unknown in Central America ; but it reappears in the northern Andean 
district of South America, where a genus (Tremarctos), distinct from Urstis, is now 
found — and this genus, oddly enough, also appears to contain the Central Asiatic 
Black or Tibetan Bear, which ranges from Baluchistan through the Himalayas to 
Japan. 

The palseontological history of the families of Central American Camivora need not 
be discussed in detail. The evidence, incomplete in details though it be, shows that 
they are without exception descended from northern forms that inhabited North America 
or Europe or both continents during Middle or early Tertiary times, the Canidse dating 
back to the Upper Eocene, the Procyonidae to the Lower Miocene in North America, 
the Ursidae to the European Upper Miocene where they apparently blend with the 



MAMMALIA. 97 

Canidse, the Mustclidse to the Lower Oligocene, and the Felidaj *, even Felis itself, to 
the Middle Miocene of Europe and North America. No extinct forms so early in 
time have yet been discovered in South America. Hence it is needless to look 
beyond North America for the imuiediate origin of the Central American and South 
American forms. But no data, palaeontological or otherwise, seem to have been 
discovered as yet to explain the almost complete absence of Canidse from the southern 
countries of Central America and the discontinuity in the distribution of the Bears 
above alluded to. Nor in the case of such genera as Galera, Grison, Nasua, and others 
is it clear whether their presence in Central America is due to northward migration 
from South America, or whether it is attributable to their settlement in the former 
area on their way to the southern continent. 

Order ARTIODACTYLA. 

The Bunodont or non-ruminant Artiodactyls are represented in America by the 
family Tayassuidse (formerly Dicotylidae), commonly known as Peccaries. The 
described species may be referred to two categories, regarded sometimes as genera, 
exemplified by the Collared Peccary {Tayassu tajacu = torquatus) and the White- 
lipped Peccary (T. pecari = labiatus). Species of the former category range from 
Texas (Guadalupe) to the Argentine, and have been recorded from Mexico (Sonora, 
Colima, Puebla), Guatemala, and Panama ; those of the second category spread from 
Campeche to Paraguay. 

Three families of ruminant Artiodactyls inhabit Central America, namely the 
Bovidge, Antilocapridae, and Cervidae. The first two, represented by the Bison {Bison), 
the Kocky Mountain Sheep (Ovis), and the Prongbuck (Antilocapra), are essentially 
North American. The Bison inhabited Mexico in historic times. The Sheep and the 
Prongbuck penetrate Mexico as far as Chihuahua. The Cervidae or Deer belong to 
two genera — Odocoileus, which extends from the United States, through Mexico to 
Panama, and thence into South America, and Mazama, ranging from Mexico (Vera 
Cruz) and Yucatan to Brazil and the Northern Argentine. 

The palaeontological history of the Peccaries is fairly well known, many genera of 
the family having been discovered in the North American Tertiaries, dating from the 
Lower Oligocene. But our knowledge of the descent of the ruminant Artiodactyla is 
singularly meagre in the matter of details. The Central American Cervidae {Odocoileus, 
Mazama) belong to a group of the family (Telemetacarpalia) which, with a few 
exceptions, is confined to America. Possibly they are descended from a form 
{Blastomeryx) which existed in North America in the Lower Miocene. Odocoileus 
itself goes back to the North American Pliocene, but it seems difficult to avoid the 

* Excluding the extinct Oligocene Machaerodonts, which have been recently claimed as their ancestors. 

BIOL. CENTK.-AMEE., Introd. Vol., June 1915. o 



98 OEIGIN, ETC., OF THE FAUNA. 

conclusion that Mazama, typically a South American genus, is a more primitive type, 
unless it be degenerate, in its simple one-spiked antlers. Nevertheless, the absence of 
extinct Deer in South America, except in late Tertiary (Pleistocene) deposits, precludes 
at present the view that the family was evolved in that country and subsequently made 
its way northwards. 

The Antilocapridae are not certainly known from deposits in North America older 
than the Pleistocene, and none have been discovered elsewhere. Possibly the North 
American mid-Miocene genus Dromomeryx was the ancestral form. 

The oldest-known forms of Caprinae (Ovis) and Bovinse {Bison) are European. 
The Bovinse do not appear to be definably distinct from the so-called Tragelaphine 
Antelopes, which in Europe go back to the Upper Miocene. A Caprine {Cridotherium) 
is of that date also. Hence it appears that the Artiodactyle Ungulates of Central 
America must be regarded as of northern descent. 

Order PERISSODACTYLA. 

Of the existing families of this order only the Tapirida; occur in Central America, 
where they are represented by two species of the genus Tapirella (ElasmognatJnis), 
namely hairdi, ranging from South Mexico to Panama, and dowi, from Guatemala to 
Nicaragua and Costa Rica. The Tapirs of South America, where they extend from 
Venezuela to the Northern Argentine, are referred to a distinct genus, Tapirus. 
Outside Tropical America the family contains but one representative, namely, lihino- 
choerus indicus, from the Malay Peninsula and Sumatra. 

Tapirs referred to a variety of genera inhabited North America from the Lower 
Eocene and Europe from the Lower Oligocene. There are gaps in the genealogy of 
the recent forms, but species akin to them have been discovered in Pliocene deposits in 
Europe and Asia and possibly in contemporaneovis beds in North America. From this 
it seems tolerably clear that the existing genera are the survivors in the East Indies 
and in Tropical America of a family formerly widely distributed in the Northern 
Hemisphere. 

Order SIRENIA. 

One genus of this order, the Manatee (Trichechvs = Manatus), occurs in the rivers 
of eastern Central America and South America. It also inhabits some of the rivers of 
Western Africa, but is not found elsewhere. 

There is evidence that the Sirenia were evolved in North Africa from the stock that 
also gave rise to the Proboscidea. Genera of Eocene age have been discovered in 
Egypt and also in the West Indies. Although Trichecus itself is only known as an 
existing genus, its presence in rivers and estuaries on the eastern and western shores 
of the Atlantic and its avoidance of the open sea have been cited as evidence of a 
continuous coast-line between Africa and Tropical America. 



MAMMALIA. 99 

■ Order EDENTATA. 

Eliminating the Old World genera Orycteropus and Manis, this order is confined to 
South and Central America. Choloepus and Bradypus, the two genera of Sloths or 
Bradypodidae, pass from South into Central America, the former as far as Costa Rica and 
the latter to Nicaragua. Myrmecophaga and Cyclopes, two of the Anteaters (Myrme- 
cophagidae), also go as far as Guatemala; the third genus, Tamanduas, extends 
northward to Tabasco and Vera Cruz in Mexico. Of the large number of South 
American genera of Armadillos (Dasypodidae) two only enter Central America, namely 
Cahassous (Lysiurus), which ranges from Brazil to Honduras, and Dasyjms { = Tatu or 
Tatusia), which overlaps it both to the north and south, with an extension from the 
Argentine to Texas, just touching the latter state at Brownsville. 

The origin and centre of evolution of the Edentata is at present unknown. The 
only extinct forms that have been claimed to be their ancestors are the Taeniodonts or 
Ganodonts of the Eocene of North America. Since this view has been authoritatively 
disputed, it needs no further consideration here. Of more importance, from our 
present point of view, is the mid-Eocene North American genus Metaclieiromys, believed 
by some to be a primitive Armadillo. It is true that the Armadillos (Dasypodidae) are 
the least specialised of all undoubted Edentates, living or extinct, and no reason can 
be alleged against the likelihood of the discovery of an early type in the Eocene of 
North America, but since Metaclieiromys unfortunately has few, if any, of the 
characters distinctive of Edentates as we know them, it cannot be maintained as 
established that the order is of North American origin. It may be suggested that 
their ancestors entered South America from Africa. However that may be, the 
evidence of fossil forms shows that South America has been the centre of evolution 
of the large number of existing and extinct genera that have been discovered. In 
South America, Armadillos (Dasypodidae) date back to the Upper Miocene, the Dasy- 
podinae {olim Tatusiinae) themselves, which are now found in Central and North 
America, ranging from that date to the Pleistocene. Hence the presence of DasypKS 
in Central America and Texas must be assigned to northward migration. And the 
same must be claimed of the Sloths (Bradypodidae) and Anteaters (Myrmecophagidae). 
No extinct forms of these families have yet been discovered, but abundant remains of a 
wholly extinct group, intermediate in many respects between the two, but less specialised 
than either, have been discovered in South American Miocene and later deposits. 
These are the Ground-Sloths (Mylodontidae and Megatheriidaj), which at a later date 
(Pliocene and Pleistocene) lived also in North America. 

Order MARSUPIALIA. 

The American Marsupials belong to two families, theDidelphyidae (or Opossums) and 
the Cienolestidae. The latter, represented by the two existing species of Cainolestes 

o2 



100 OEIGIX, ETC., OF THE FAUNA. 

from Ecuador and Colombia, does not appear to reach Central America ; but no fewer 
than five genera of the former occur in our area, namely Didelphys, Metachirus, 
Caluromys [= Philander), Marmosa, and Chironectes. Of these JJidelphys is the only 
one that is found in North America, where it ranges from about the 40th parallel 
of N. lat. to Texas, thence passing southwards through Mexico, Nicaragua, Yucatan, 
Guatemala, and Panama as far as the Argentine and Chili. The others are Central 
and South American, Marmosa ranging from Mexico (Oaxaca), Costa Rica, and Panama 
to Chili, Metachirus from Mexico (Vera Cruz), Guatemala, and Costa Rica to the 
Argentine, Caluromys from Mexico (Tabasco) to Paraguay, and Chironectes, the 
aberrant water-opossum, from Guatemala to Brazil. 

Extinct Marsupials belonging to the Didelphyidae, and closely related to the existing 
genera of that family, have been discovered in Lower Eocene deposits in North 
America and as late as the Lower Miocene in Europe, and in what are believed to be 
Upper Cretaceous beds in South America. Formerly, therefore, the family was very 
widely distributed. Nevertheless, Opossums appear to have survived uninterruptedly in 
South America, at all events since the earliest Tertiary times, and that country is now 
tlieir headquarters. These facts, coupled with the absence of palteontological evidence 
that the group survived in North America after the Oligocene and in Europe after the 
Miocene, justify the supposition that the existing genera are recent immigrants from 
South into Central and North America. 

SUMMARY AND CONCLUSION. 

Origins of the Central American Mammalian Fauna. 

The foregoing account shows that the Mammalian fauna of Central America is a 
mixture of two elements, namely, of forms which are dominant in North and South 
America respectively ; and the palseontological history, during Tertiary times, of the 
orders concerned, supplies — in part, at all events — the explanation of the intermixture. 
So far as this history has been read, it furnishes strong evidence that the Mammals 
may be referred to two categories. To the first belong the Insectivora, Carnivora, 
Artiodactyla, Perissodactyla, and the Rodentia (with the exception of the Hystrico- 
morphs and possibly some of the Cricetine Myomorphs), which were evolved 
through long ages in the Northern Hemisphere and inferentially passed from North 
into South America by way of Mexico and Panama. The second comprises the 
Edentates, Primates, Marsupials, and the Hystricomorphous Rodents which, from 
whatever country they may originally have come, have undergone a prolonged course 
of evolution in South America and migrated thence into Central or even North 
America. That is to say, whereas North and South America have been independent 
geographical centres for the evolution of Mammals in the Western Hemisphere, the 
part played in the main by Central America has been that of a bridge joining these 
centres together and permitting the intermingling of the independently developed 



MAMMALIA. 101 

faunas. A few genera of Chiropteia and Rodentia restricted, so far as is known, to 
Central America, may have been evolved within the area ; but these do not alter the 
general character of the country as a faunistically transitional tract between the 
northern and southern portions of the Western Hemisphere. 

That the intermingling and cross-migrations, which give the stamp to the existing 
Central American Mammalian fauna, began in the Miocene and have continued 
uninterruptedly since that date may be inferred from palseontological and geological 
data, which support the conclusion that the Northern and Southern Americas, separated 
by sea over what is now the isthmus of Panama from the earliest Eocene (Paleocene) 
to the end of the Oligocene, were finally joined by the elevation of that isthmus during 
the Miocene Period. 

Possibly, as held by some authorities, Central America to the north of Costa Rica 
and Panama was temporarily connected with South America by way of the Greater 
and Lesser Antilles in the Early Oligocene ; and possibly there was a still earlier union 
during the Cretaceous of the western portions of North, Central, and South America. 
However that may be, it seems tolerably certain that the main streams of migration 
passed by way of Costa Rica and Panama in comparatively recent Tertiary times. 

Reference has more than once been made to the possibility of South America 
having received the ancestors of some of its characteristic forms of Mammals (Platy- 
rhine Monkeys, Hystricomorph Rodents, Sirenians) from Africa by a direct trans- 
Atlantic bridge between the two continents. Other groups of animals supply evidence 
for the existence of this union. But if the mammals made use of it, it must have 
endured into early Tertiary times. This, however, has been disputed. 

Another point, also under discussion, may here be referred to, although it has no 
direct bearing, so far as is known, upon the fauna of Central America. This is the 
possibility of a direct connection between South America and Australia. Perhaps the 
most cogent evidence for this is supplied by certain genera of invertebrates, probably 
in great part ancient types. So far as Mammals are concerned, the evidence rests 
upon the claimed relationship between the so-called Tasmanian or Marsupial Wolf 
(Ihylacinus) and some extinct (Upper Miocene) Sparassodont Mammals of South 
America, coupled with the later date of fossil Marsupials in Australia, the absence of 
their remains in Tertiary deposits in China, and the absence of living forms in South- 
east Asia to the west of " Wallace's Line." Also there is the undeniable kinship 
between the Australian Dasyurids and the American Didelphyidae, of which Marmosa 
is alleged to be the most primitive type. It would be out of place further to discuss 
these questions here ; but the facts, as they stand, are suggestive of the origin of the 
Australian Marsupials from a South American stock. Moreover, if Thylacinua be of 
the same family as the Sparassodont Prothylacinus, it may be held as perhaps 
probable that the connecting land-mass permitting the migration persisted into early 
Tertiary times. 



102 OEIGIN, ETC., OF THE FAUNA. 

Faunistic Divisions of Central America. 

Central America has been described above as transitional with respect to its 
Mammalian fauna between North and South America. This statement, however, is 
perhaps too general and apt to give a mistaken impression of the actual facts ; for the 
transition cannot be described as complete. Its incompleteness, however, is not due 
to the existence of any physical barriers to migration, unless temperature and moisture 
can be described as such. But, as will appear in the sequel, the faunistic do not 
coincide with the political divisions of this land-area. 

The central portion of Mexico forms an extensive plateau rising some 9000 feet 
above the sea, with mountains nearly twice that altitude. This tableland to the north 
is continuous with, and closely resembles in its characters, a great tract of comparatively 
dry territory stretching into North America and embracing the southern parts of 
California, Lower California, Nevada, Arizona, and nearly the whole of Texas. 
Southwards the plateau is continued by the mountain ranges which stretch throughout 
the southern States of Central America In Mexico it rises somewhat abruptly 
from the lower-lying country towards the coast on each side of it; and this country, 
covered for the most part with tropical forest, extends to east and west of the plateau 
up to or almost up to the confines of the United States. Tropical forest-conditions 
also prevail over the greater part of Central America to the south of the plateau, 
although, as has been stated, the country is almost everywhere broken up by 
mountains rising several thousand feet above sea-level. 

It is needless to recapitulate here the facts set forth above regarding the distribution 
of the families and genera of Mammals inhabiting Central America. It is clear, how- 
ever, that those believed to have undergone a long course of evolution in South 
America (Primates, Edentates, Hystiicomorphous Rodents, and Marsupials) are wholly, 
or in the main, restricted to the forested tracts above described, although not by any 
means occurring throughout such districts. Most of the genera and famiUes do not 
pass north of Vera Cruz, in Mexico, and Guatemala. Notable exceptions are the 
Armadillo (Dasi/pus) which reaches Texas, some of the Opossums which occur on the 
Mexican plateau — one, indeed, penetrating far to the north in the United States, — and 
the Porcupine [Erethizon), belonging to the same family as Coendou and ranging 
from Arizona to Alaska. 

Similarly, with the groups which, although known to have been evolved in North 
America or Eurasia, are now dominant in the southern tropical districts of America, 
such as the Tapirs, Peccaries, Brockets (Mazama), most of the FelidEC, some Procyo- 
nidse {Nasua, Potos), and Mustelidse [Tayra, Grison). These genera, usually regarded 
as intruders from the south, though the evidence on that head seems inconclusive, are 
mostly restricted to forest-covered tracts of Central America. And the same thing 
applies to the Bats of the families Phyllostomidas, Desmodontida;, etc. 



MAMMALIA. 103 

On the other hand, there are a great many genera like the Prongbuck {Antilocapra), 
Bison (formerly), Rocky Mountain Sheep, Beaver, Prairie-Marmot {Cynomys), Chipmunk 
[Eutamias), Black Bear, Wolf, Lynx, and other typically North American forms which 
are restricted in Central America to the Mexican plateau or to the highlands just to 
the south of it. 

These differences in distribution exist quite apart from the occurrence throughout 
Central America of many families and genera like the Hares (Leporidae), the typical 
Squirrels {Sciurus etc.), the Shrews (Soricidse), Pocket Gophers (Geomyidae), Raccoon 
{Frocyon), Deer (Odocoileus), and others. 

Zoogeographers have given practical expression to the facts above set forth by 
referring the plateau of Mexico to the same zoological region as the southern States of 
North America. Out of a number of titles that have been proposed for this region, 
Sonoran is the one preferred. The rest of Central America, on the other hand, is 
regarded as a suhregion of the Neotropical Region — sometimes called the Neogaeic 
Realm, — which embraces, in addition, the Antilles and the whole of South America. 
On the western side of the plateau this region stretches to about lat. 25° N., and on 
the eastern side a little farther, stopping short near the Nueces River in Texas. 

The subregional distinctions between the tropical portions of Central America and the 
adjacent portions of South America are not sharply defined, and rest upon the not very 
satisfactorily established statistics of the comparatively small number of typically 
South American genera which occur in those areas of Central America, and the 
presence in the latter of certain Sonoran forms (Soricidae, Geomyidae) whose southward 
range practically stops short at Panama. 

Similarly, the Antillean subregion is characterised mainly by the poverty of its fauna 
in types occurring in Central and South America. It is noticeable that the orders of 
known northern origin are practically absent. Some of the Chiroptera (Eptesiciis, 
Myotis) are, perhaps not surprisingly, exceptional. Most of the genera of this order 
belong to the Tropical American family Phyllostomidae. For the rest, the Hystrico- 
morpli Rodents are represented by Capromys, Plagiodontia, and Loncheres, all Octodonts, 
the first two being peculiar to the subregion, by Basyprocta, and possibly by a species 
of Coendou. Very interesting is the occurrence of a Cricetine Myomorph, Megalomys, 
peculiar to the Lesser Antilles. Monkeys, however, are absent, and the Edentates are 
represented only by one species of Armadillo {Dasypus) from Grenada. Finally, the 
Insectivora are exemplified by the Cuban and Haitian Solenodon, whose nearest living 
allies are the Centetida) of Madagascar {Centetes etc.) and West Africa {Potamogale). 
The family Solenodontidae, however, dates back to the Lower Oligocene in North 
America, where one genus, Microp)ternodus, has been discovered. 

Until the Mammalian Palaeontology of the West Indies has been worked out, the 
geological history of these islands cannot be accurately read. Nevertheless, temporary 
union between the Lesser Antilles and South America is indicated by the occurrence 



104 OEIGIN, ETC., OF THE FAUNA. 

of such genera as Loncheres, Basyprocta, and Dasypus. Furthermore, the restriction of 
Capromys to Cuba, Jamaica, and some of the islands towards Central America and 
the kinship between this genus and living and fossil South American genera suggest 
a connection between Central America and those islands of the Greater Antilles. 
But this conclusion cannot at present be reconciled with the absence of other Centrul 
American forms, both of northern and southern origin, from the Greater Antilles. 
If, on the other hand, the Hystricomorphs passed into South America from Africa or 
South Europe, and if the West Indies formed part of the transatlantic land, the 
faunistic resemblance between Haiti, Cuba, Jamaica, and South America supplied by 
Capromys and Plagiodontia may be due to the derivation of the fauna from a common 
African source. The relationship between Solenodon of Cuba and Haiti and the 
Afro-Mascarene Centetidte has an interesting bearing on this question, although, if 
Micropternodus be, as alleged, a Selenodont Insectivore, Cuba and Haiti must 
presumably have been connected with some part of North America. 



PRINCIPAL WORKS CONSULTED. 

Alston, E. R. — Biologia Centrali- Americana. Mammalia, 1879-1882. 

Gadow, H. — The Wanderings of Animals. Cambridge Manuals of Science and Literature, 1913. 
Lydekker, R. — A Geographical History of Mammals. Cambridge Geographical Series, 1896. 
Mekriam, C. Hart. — " The Geographical Distribution of Life in North America, etc.," Proc. 
Biol. Soc. Wash. vii. pp. 1-64, 1892. 

Miller, G. S. — " List of North American Land Mammals, etc.," Bull. U.S. Nat. Mus. 1912. 
OsBORN, H. F. — The Age of Mammals. Macmilian & Co., New York, 1910. 

ScHARFF, R. P. — Distribution and Origin of Life in North America. Constable & Co., London, 

1911. 
Scott, W. B. — A History of Land Mammals of the Western Hemisphere. Macmilian & Co., 

New York, 1913. 
Trouessart, E. L.— Cat. Mamm., Suppl. 1899-1904. 



[ 105 ] 

REPTILIA, BATRACHIA, AND PISCES. 

By C. Tate Regan, M.A. 

It has been found convenient to deal with these groups in the reverse order to that 
indicated in the heading of the chapter, and to take the Fishes first. 

PISCES. 

Marine Fishes.— It is well known that the fishes of the Pacific coast of America, 
from California to Peru, are quite distinct from those of the tropical Indo-Pacific, and 
are related to those of the Antillean district. At one time it was thought that a large 
proportion of the species were found on both the Atlantic and Pacific coasts, but the 
most recent work tends to show that when pelagic fishes of wide distribution are 
eliminated the number of species identical on the two coasts is very small. There 
are, however, many closely related species that represent each other on the Atlantic 
and Pacific coasts, and the inference is that each pair has evolved from a parent form 
that existed when the two oceans were connected. There is evidence that North and 
South America were separated by sea during the Eocene, and became one continent 
in the Miocene ; if this were so, the similarity and degree of dissimilarity between 
the tropical shore-fishes of the Atlantic and Pacific coasts would be satisfactorily 
explained. 

Fresh- water Fishes. — Fishes that are found in lakes and rivers sometimes belona: 
to marine species that enter fresh water for purposes of breeding or feeding ; such 
species may form permanent fresh-water colonies or races, and these lead to fresh-water 
species of marine genera and fresh-water genera of marine families. These are 
unimportant in zoo-geography, but there are many families and even some orders that 
are confined to fresh water, and appear to have evolved their genera and species in 
fresh water ; their dispersal has depended on hydrographical changes, such as the union 
of rivers formerly distinct or the capture by one stream of the tributaries of another, 
and for most of them the sea appears to be an impassable barrier. Such fishes may be 
termed true fresh-water fishes, and they are of the highest importance as indicating 
former land-connections or ancient lines of severance. 

The Neotropical Region.— South America has a rich and varied fish-fauna, 
surpassing that of any other region for wealth and individuality. As in other parts 
of the world, except the Australian Region, the majority of the true fresh-water fishes 
belong to the order Ostariophysi *. This order comprises two well-marked suborders, 

* Cf. Ann. & Mag, Nat. Hist. (8) viii. 1911, pp. 13-32, 553-577. 

BIOL. CENTB.-AMER., lutrod. Vol., June 1915, p 



106 OKIGIN, ETC., or THE FAUNA. 

Cyprinoidea (Chaiacins, Electric Eels, Carps, Loaches) and Siluroidea (Catfislies). 
Of the Cyprinoids the Cypriniformes are absent from South America ; the Characi- 
formes (Characins) are represented by five families — four endemic and the other, the 
most generalized, found in Africa also; whilst the Gymnotiformes (Electric Eels) are 
peculiar to the Neotropical Region. The Siluroids include the archaic Biplomystes 
of Chile and eight endemic families, of which the most generalized, Pimelodidae, is 
related to the African and Indian Bagridse. The only other Neotropical fishes of much 
importance geographically are the Cichlid Perches, also found in Africa, for the 
Cyprinodonts are partly marine and the sea has aided in their dispersal. 

Origin of Neotropical Pishes.— it has been suggested by various authors that 
the ancestors of the characteristic neotropical faunal groups may have reached South 
America in late mesozoic or very early tertiary times either from Australia via 
Antarctica, from North America, or from Africa via an xltlantic continent. Recently * 
I have attempted to show that the evidence for the supposed connection with Australia 
does not warrant the conclusions that have been reached ; so far as the true fresh-water 
fishes are concerned, there is only one family common to Australia and South America, 
the Osteoglossidpe, a generalized, ancient, and widely distributed group represented at 
the present day by a few remnants. As to North America, it has not, at the present 
day, a single family in common with South America, and the fossils show that it has 
been just as distinct throughout the tertiary. Cat-fish {Rhineastes) , an Osteoglossid 
{Dapedoglossus), and a Perch [Priscacara) from the Green River Shales (Lower Eocene) 
iu Wyoming have been cited as showing neotropical affinities f. In my opinion, 
Rhineastes may belong to the cosmopolitan and marine family Ariidfe, Dapedoglossus 
seems to be nearer to the Indo-Australian Scleropages than to the South American 
Osteoglossum, and Priscacara is not a Cichlid. All the Cichlidae, whether African or 
American, have the palate toothless, and the caudal fin formed of 16 principal rays, 

14 branched. Priscacara has teeth on the vomer and 17 principal caudal rays, 

15 of them branched. After examination of the specimens in the British Museum, 
I conclude that Priscacara is a member of the endemic Nearctic family Centrar- 
chidaj, and is closely related to the modern Pupomotis, in which genus the 
enlargement and coalescence of the lower pharyngeals may also be seen. If the 
ancestors of the present Neotropical fishes came from North America, no trace of 
them has yet been discovered. 

The hypothesis of a former land-connection between South America and Africa 
receives strong support from the Fishes ; although no genera of the Characidae and 
Cichlidse are common to the two continents, the close relationship of Rrycon and 

• Brif. Antarctic 'Terra Nova' Exped., Fish. 1914. 
t Osborn, ' The Age of Mammals,' p. 13G. 



EEPTILIA, BATRACHIA, AND PISCES. 107 

Alestes of the Characidse, and of Acara and Paratilapia of the Cichlidae, seems 
apparent. Further support is derived from the Lepidosirenidae, with Lepidosiren in 
South America and Protopterus in Africa, and from the relationship of the Pimelodidae 
and Bagridse. 

Evolution and Dispersal of Neotropical Fishes,— Since all the genera and 

most of the families of true fresh-water fislies of the Neotropical Eegion are peculiar, 
and the majority of the endemic families may be regarded as specialized Characidge 
or PimelodidiE, it may be inferred that the present fish-fauna has evolved in South 
America from a few ancestral types, and that it has received no immigrants from other 
regions since the earliest of the Tertiary. 

It is generally accepted that the Antillean Region was submerged during the Eocene, 
and that the invasion of Central America from the south dates back not earlier than 
the Miocene. The distribution of the fishes is in harmony with this supposition, for, 
Avhilst most of the Neotropical families have a wide range in South America, only four 
of them extend north of the isthmus of Panama. One of these is the Gymnotidse, 
represented by Gymnotus carapo {Giton fasciatus), found everywhere from Montevideo 
to Guatemala ; the others are the Pimelodidae, Characidae, and Cichlidae. The 
Pimelodidae are represented mainly by about twenty species of the large and widely 
distributed South American genus Bhamdia, which ranges north on the Atlantic side 
to southern Vera Cruz, but has not reached the Balsas nor the Mexican Plateau. 

A few genera of the Characidae have got as far as Bhamdia, and it is only two or 
three forms that are scarcely specifically distinct from Tetragonopterus {Astyanax) 
ruiilus, found everywhere in South America north of the La Plata, that extend the 
range of this family to the Balsas and on the Atlantic slope to the Rio Grande. 

In the Characidae and Pimelodidae invasion of Central America has resulted in the 
differentiation of a number of species, but has produced no types that are markedly 
distinct from their relatives in South America. 

The Cichlidae are in many respects peculiar ; the Central and South American 
species are about equal in number and for the most part are generically or at least 
subgenerically distinct, so that it is usually possible to recognize at a glance whether 
a species is Central or South American. The South American types are the more 
generalized, for the majority have three anal spines and simple conical teeth ; in the 
Central American genera the number of anal spines is increased and various 
specializations of the dentition occur. The great lakes of Nicaragua have a highly 
specialized endemic Cichlid fauna; many species are found in the region between 
Panama and southern Vera Cruz, and a few in the Atlantic coast-streams northwards 
to the Rio Grande. Only one species, belonging to the genus or subgenus Parapetenia, 
is found in the Balsas and a related form in the lowland streams of Sinaloa ; a species 
of this type also occurs in Cuba. The Central American Cichlidae are a difficult group 

p2 



108 OlilGlN, ETC., OF THE FAUNA. 

for the systematist ; the species are closely related and the genera ill-defined. They 
give the impression that their number and variety, as compared with the Characidae 
and the Pimelodidee of this area, are not due to an earlier migration, but to a greater 
capacity for differentiation and a more rapid evolution. 

The Nearctic Kegion.— North America has a fish-fauna as different as possible 
from that of South America. The Ostariophysi are represented by the Catostomidse 
(Suckers) and by a family of Cat-fishes, the Amiuridse, each restricted to this region 
except for one or two species in China ; in addition, there are a number of Leuciscine 
Cyprinidse, a group well represented in Eurasia. The Perches belong to the endemic 
family Centrarchidse and to the Percidae. The Cyprinidse, Percidse, Esocidse, and 
Umbridae are holarctic, but there are several endemic families, Hiodontida;, Per- 
copsidse, etc., in addition to those mentioned above, that make this region quite 
distinct from the Palsearctic. 

Origin of Nearctic Fishes.— Numerous fresh-water fishes have been described 
from early Tertiary deposits in North America. Lepidosteidaj and Amiidae were 
already present in the Upper Cretaceous or Basal Eocene of North America and 
Europe. An interesting assemblage is known from the Lower Eocene (Green Eiver 
shales) of Wyoming, including Percopsidse or Aphredoderidae (Anrphilaga and Erisma- 
topterus), Percidae {Mioploms), and Centrarchidae {Priscacara), as well as the Osteo- 
glossid Dapedoglossus ; there are some other fishes that may have been marine as well 
as fresh-water, e. g. Xiphotrygon (Trygonidae), Notogenus (Chanidae), Biplomystus 
(Clupeidae), and Rhineasftes (Ariidae). Catostomidae are also known from the Eocene 
and Amiurus from the Lower Miocene. 

The known history of the Nearctic fishes is in harmony with the supposition that 
North America separated from Eurasia at the end of the Cretaceous and developed 
several of its endemic types during the Eocene, and that since the Eocene one or 
more connections with Eurasia have brought Leuciscine Cyprinidae to America, and 
have established two or three Catostomids and Amiurids in China. The identity cf 
some of the more northern species of the two continents shows that a connection 
across the Behring Sea must have persisted until a recent date. 

Dispersal of the Nearctic Fishes. — In the Mississippi all the characteristic 
nearctic types are represented ; to the north, west, or south there is a marked 
impoverishment. In the Eio Grande, Hiodontidae, Percopsidae, Aphredoderidae, 
Umbridac, and Esocidae are absent, and the Percidae and Centrarchidae are reduced to 
a few species only ; there are several Cyprinidae and Catostomidae. Isolated streams 
and lakes of Chihuahua and Durango, the portions of the Yaqui and Mezquital Rivers 
east of the Sierra Madre, and the rivers of Tamaulipas and northern Vera Cruz have a 
fish-fauna essentially similar to that of the Eio Grande. On the southern part of the 
Mexican Plateau, the Eio Grande de Santiago or Lerma above the falls, with the 



REPTILIA, BATRACHIA, AND PISCES. 109 

isolated lakes of the states of Michoacan and Mexico, constitute an important system 
with a very characteristic fish-fauna formed almost entirely of endemic species. Only 
three families of Nearctic fishes are present ; the Catostomidse are represented by a 
single species of Moxostoma and the Amiuridac by an Amiurus, Avhilst there are 
a dozen Cyprinidfe, nearly all belonging to endemic genera. The majority of the 
fishes belong to two groups, each with about a score of species, and each belonging to 
a family that includes a number of marine fishes that enter fresh water. These two 
groups are characteristic of the Lerma System and barely overstep its limits ; one is 
the somewhat heterogeneous Atherinid genus Chirostoma, the other is the Cyprinodont 
subfamily Characodontinse, which includes five well-marked genera, and diff"ers from 
the Fundulinse in that its species are viviparous. 

The Lerma System has evidently long been isolated, and has been a centre of 
evolution. The chain of volcanoes that border the Mexican plateau to the south have 
barred invasion on that side, and from the north only a few nearctic types have 
reached the Lerma. The main elements of its fish-fauna, Chirostoma and the Chara- 
codontinae, have in all probability evolved each from a single ancestral type that 
entered from the sea, from which it is now separated by inaccessible falls. 

The Kio Balsas lies to the south of the Mexican plateau, and is a large river that 
flows into the Pacific ; it has not been thoroughly investigated, but appears to have 
a comparatively poor fish-fauna. In addition to a Lerma type (Goodea) that has got 
into the headwaters of its northern tributaries, there is a Cat-fish (Amiurus) and a 
Cyprinoid [Notropis) ; two neotropical species balance these. The high mountains 
that bound the Balsas System have evidently rendered immigration a difficult feat. 

It is in the lowlands of the Atlantic slope that conditions have been more favourable 
to migration. Lepidosteus, absent from the Mexican plateau and from the Balsas, has 
reached Panama, and a species of Ictiobus (Catostomidse) and an Amiurus occur in 
the Usumacinta, balancing the Cichlid and Characid species of the Rio Grande. 

Dispersal of Neotropical and Nearctic Tishes compared. — It has been 

shown above that comparatively few elements of the rich neotropical fish-fauna extend 
north of the Isthmus of Panama, and that of these only the CichlidiE exhibit much 
diversity and specialization. In Central America and in rivers of the Atlantic slope 
of Mexico south of Vera Cruz the fish-fauna is almost exclusively neotropical, but 
only two neotropical types have reached the Balsas, none have surmounted the volcanic 
chain that borders the Mexican plateau, and only half a dozen have penetrated north 
of Vera Cruz in the Atlantic coast-streams of northern Mexico. In these streams 
there are many nearctic species, but in Atlantic rivers south of Vera Cruz only three. 
The Rio Grande exhibits a paucity of nearctic types as compared with the Mississippi, 
and this is still more marked in the Lerma System, which possesses a peculiar endemic 
fish-fauna. In the Balsas, equally isolated from the north and the south, nearctic and 
neotropical species balance, but there are only two or three of each. 



110 OEIGIN, ETC., OF THE FAUNA. 

It is probable that the distinctness of the North and South American faunas is 
more marked, and that where they meet there is less overlapping in the case of the 
fresh-water fishes than in any other group of animals. 

Fresh-water Fishes.— General Conclusions.— The South American fresh-water 

fish-fauna is as distinct as the Mammalian fauna would have been if a large part of 
the endemic fauna had not died out in late Tertiary times, and been replaced by the 
invasion of types that had evolved elsewhere. 

Turning to the other parts of the world, we find that the distinctness of the 
Australian Region, marked in the Mammalia, is emphasized by the Fishes. The 
effectiveness of the sea as a barrier is strikingly illustrated by a comparison between 
Borneo, with its hundreds of species of Cyprinoids, Siluroids, Anabantoids, etc., and 
Celebes, without a single indigenous true fresh-water fish. The other four regions are 
well characterized, although less distinct, for the Indian has certain relationships with 
the Ethiopian and Palsearctic, and the last with the Nearctic ; there has been a certain 
interchange, but in each case the endemic groups preponderate. 

The general impression derived from a study of the fresh-water fishes of the world is 
that many of the families were in existence at the beginning of the Tertiary, that for the 
most part they have evolved in the areas they now occupy, have dispersed slowly, and 
have never had a distribution much wider than at present. For Mammals, it may be gene- 
rally stated that they have evolved rapidly and spread rapidly, and have found it easier 
to reach a country than to live in it. For fresh-water fishes migration has been much 
more difficult, survival relatively easy ; hence their great importance in zoogeography. 

BATRACHIA. 

BatrachianS. — These resemble fresh-water fishes in their inability to swim across 
the sea, and their eggs are no more likely to be transported over the sea than those of 
fresh-water fishes. But it is probable that accidental transmission has played a part in 
the dispersal of some arboreal and terrestrial frogs ; at any rate, this seems the most 
reasonable explanation of the distribution of certain genera and species in the Indo- 
Australian Archipelago. Even the most aquatic types can travel overland from 
one stream to another, and consequently are able to migrate rapidly when conditions 
are favourable. 

Nearctic and Neotropical BatrachianS.— The Batrachian fauna; of these two 
regions are less distinct than the fish-faunae, but are, nevertheless, very different. 
Urodeles are scarce in South America, Ccecilians are absent from North America ; the 
majority of the Neotropical Frogs are Cystignathidge and Hylida;, families but sparingly 
represented in the Nearctic Region. 

The mountains that fringe the Mexican Plateau form the boundary between the 



EEPTILIA, BATRACHIA, AND PISCES. Ill 

the Nearctic and Neotropical regions ; only a small proportion of the tropical genera 
have gained a footing on the plateau, and but a few undoubted northerners have 
spread far to the south beyond it. 

The Urodela are almost exclusively holarctic and aquatic ; except the monotypic 
Thorius, known only from the mountains of Orizaba and Oaxaca, but two genera, Ambly- 
stoma and Spelerpes, extend southwards into Mexico and Central America. Each of 
these genera has several species in the United States, and whilst Amblystoma has one 
in the mountains of Siam, Spelerpes includes one from Sardinia and Italy. These 
isolated species suggest that both genera had formerly a v/ider and a more northerly 
range, and that only one Old World species of each managed to survive the glacial 
epoch by migrating southwards. In America Amblystoma extends on to the Mexican 
plateau and the mountains that fringe it, but Spelerpes includes about seven species 
from the United States, fifteen from the mountains of southern Mexico and Central 
America, three from the Andes of Peru and Colombia, and one from Haiti. It seems 
evident that this genus began to spread southward as soon as the elevation of Central 
America enabled it to find a congenial temperature at high altitudes. 

The CoBcilians are vermiform, apodal, burrowing animals that inhabit the Neo- 
tropical, Ethiopian, and Indian regions. There are a few Central American species, 
and one of these, Dermophis mexicanus, ranges north to southern Vera Cruz. Nothing 
is known as to the past history of this group, and they may or may not be ancient 
inhabitants of South America. 

The Anura, or Frogs and Toads, may be aquatic, terrestrial, arboreal, or fossorial. 
Some genera and species have an extremely wide range, and, although it is evident 
that the sea forms an impassable barrier for the majority, it seems probable that 
accidental transmission has played a part in the dispersal of genera such as Bana 
and Hyla. 

The two main divisions of the Anura — Firmisternia and Arcifera — may or may 
not be natural, but the further classification into families is unsatisfactory ; the 
CystignathidtE, for example, are heterogeneous, as the Australian genera are very 
distinct from those inhabiting South America, the latter appearing to form a natural 
group. The Bufonida) also may be an artificial assemblage. 

To the Firmisternia belong the Ranidae and Engystomatidse, abundant in South 
America, Africa, India, and ranging eastwards to New Guinea and beyond. There 
are several endemic South American genera, and two or three v^ith Central American 
species also. Engystoma extends from Paraguay to Texas and Florida. Ilypopachus 
has one species ranging from Brazil to Costa Rica, another from Ecuador to 
Guatemala, and a third from Western Mexico to Texas. The only other genus of 
the group found in North America is the nearly cosmopolitan JRana, which extends 



112 OEIGm, ETC., OF THE FAUNA. 

southward to the Andeau region of South America, and is doubtless a recent 
immigrant. Gadow suggests that on reaching South America it gave rise to the 
endemic neotropical genera referred to the Ranidse, but it is possible that these are 
derived from an ancient neotropical stock, for at least one recent genus of the Ranidae 
dates back to the Eocene, and the family itself may have a greater antiquity. 

Of the Arcifera, the American Cystignathidse are almost exclusively neotropical ; 
there are several endemic South American genera, and of those that range northwards 
into Central America only Syrrhophus, Ilylodes, Leptodactylus, and Borborocoetes are 
represented on the Mexican plateau by a few species ; none range further north. 

The Hylidae are mainly neotropical and arboreal, and except for one or two species 
of Ilyla are absent from the Mexican plateau. Only Hyla ranges outside this region 
through North America to temperate Eurasia and southwards through China to 
Burma, then comes a gap, for the Australian Region is rich in species, only one of 
which crosses Wallace's Line into the Sunda Islands. The presence of Hyla arhorea 
in Madeira and the Canaries, the range of //. dolichopsis from Sumatra to the 
Solomons, suggest that these tree-frogs may be carried across the sea, and that their 
wide distribution may be partly due to this ; their absence from Africa and India is 
remarkable, but there are other equally curious cases of discontinuous distribution 
in genera of no great antiquity, and it is unnecessary to infer some special connection 
between Australia and South America to explain this one ; nor does the distribution 
of the Cystignathidae support this view, for the Australian members of the family are 
by no means closely related to the American ones, and their common ancestor must 
date back well into mesozoic times. 

Of the American genera referred to the Bufonidse two are strictly neotropical, 
Engystomops ranging from South America to Tehuantepec, and the monotypic 
BJdnophryne occurring on the Atlantic slope from Vera Cruz to Guatemala. The 
third genus, Bufo, is cosmopolitan except for the Australian Region ; there are a 
number of species in Central America and southern Mexico, whilst others on the 
Mexican Plateau range into the southern United States. Except for two species in 
Celebes, Bufo is absent from the Australian Region, although there are many species 
in the Malay Peninsula and Archipelago ; this is exactly the reverse of what occurs in 
Hyla, and Bana differs from both, as it is well represented in both Indo-Malayasia 
and Austro-Malayasia eastwards to the Caroline Islands. 

Scaphiopus, the American representative of the family Pelobatidse, is related to the 
European Pelobates ; there are about eight species from the United States and the 
Mexican Plateau ; these are diggers in the sand, and do not range southwards beyond 
the mountains of Southern Mexico. 

Dispersal of Batrachians. — The Anura and Urodela are both known to date 
back to the Jurassic, and this, as well as the presence in New Zealand of a Toad 



EEPTILIA, BATRACHIA, AND PISCES. 113 

(Liopelma), indicates tliat in all probability some of the Batrachian families are more 
ancient than those of fresh-water fishes. There seems no reason to doubt that they 
can and have spread rapidly, but probably their main distribution was accomplished 
before the Tertiary. The Urodela have in all probability always been northerners, 
and have only recently penetrated into South America. The Ccecilians are tropical, 
but their absence from the Australian Region rather tells against the idea that they 
are a very ancient group. Many families of the Anura are widely distributed 
and probably ancient ; the Cystignathidae and Hylidae seem to be old neotropical 
families. 

EEPTILIA. 

Reptiles. — Leaving out the aquatic members of this group, some Reptiles can and 
do swim across the sea ; others that are arboreal or that burrow into or hide in trunks 
of trees are peculiarly liable to accidental transmission over the ocean. Nearly all 
can travel quickly overland, so that in all probability the dispersal of many groups 
has been accomplished rapidly. The diiference between the Nearctic and Neotropical 
regions is decidedly less marked than in the Batrachians, but the boundary is 
the same. 

Of the groups of Reptiles represented in Central America the Crocodiles and 
Tortoises are ancient and seem to have been cosmopolitan at the beginning of the 
Tertiary ; a study of their present distribution leads to no very important results. 
The Lizards and Snakes are the dominant orders of the present day, but their past 
history is almost unknown ; in all probability the principal families date back to early 
Eocene times, and as they developed during the Tertiary they may have used the same 
routes of migration as the Mammalia. 

Dr. Gadow's papers on the Reptiles and Batrachians of Mexico and Central 
America have been referred to above (p. 59) and are of great interest, especially 
when they deal with the effects of altitude, climate, terrain, etc., on the distribution ; 
his deductions as to the origin and migrations of the different groups are necessarily 
based mainly on their present distribution, and are to that extent unsatisfactory. The 
Mammals, whose evolution during the Tertiary, extended migrations, survivals, and 
extinctions are very thoroughly known for several orders, teach that it is very unsafe 
to deduce centres of origin and dispersal from present distribution alone. 

Lizards. — There appear to be no families of Lizards restricted to South America, 
but most of the 35 genera of one family, the Teiidae, are peculiar to that continent ; 
half a dozen South American genera range into Central America, two reaching 
southern Mexico, and one — Cneiuidojihorus — the United States. This genus includes 
species on the Mexican plateau, as well as in the lowland forests. The conclusion 
that this is an old neotropical family seems not unreasonable, but may be incorrect ; 
the neotropical Monkeys furnish a parallel. 

BIOL. CENTR.-AMER., Introd. Vol., June 1915. q 



114 OKIGIN, ETC., or THE EAUNA. 

The family Amphisbajnidse comprises burrowing Lizards, with limbs reduced or 
absent ; the majority of the genera are neotropical or African, but the most 
generalized genus, and the only one with fore limbs, Chirotes, is found in California 
and Mexico, and the least specialized of the rest, Blanus, inhabits the countries round 
the Mediterranean. Moreover, Rhineura of Florida is known to date back to the 
Oligocene of Dakota. Here an American distribution similar to that of the Teiidse 
is coupled with indications that the group may have been originally a northern one 
that migrated southwards into Africa and South America. 

The Scincidse, widely distributed in the Old World, have in all probability reached 
America quite recently, as the three genera found in America are also Asiatic. 
Eumeces has several North American species, and extends southward over the plateau to 
southern Mexico. Lygosoma laterale of the southern United States ranges southward 
to southern Mexico, and in Guatemala is replaced by the closely related L. assatum. 
L. laterale is almost identical with L. reevesi of China and Burma, and it may be that 
in the warmer climate of Pliocene times the ancestral type ranged northward to the 
land connecting Asia with America. The third American genus, Mahuia, is found 
also in Africa and southern Asia; there is an Antillean species, one from Costa Rica, 
and four from South America, one of these extending northward to Yucatan and 
southern Vera Cruz. This neotropical distribution of a genus that in all probability 
came through North America parallels that of Tapirus. 

The Iguanidse are American, except for two genera in Madagascar and one in the 
Friendly and Fiji Islands. They are also known from the Eocene of Europe, and are 
evidently an ancient group of former wide distribution ; the presence of two peculiar 
genera in the Galapagos suggests their antiquity in South America. The South 
American genera are numerous ; the majority are restricted to South America, but 
some are also Antillean, and some extend through Central America as far as southern 
Vera Cruz ; there are also two or three genera peculiar to Central America. There 
are about ten genera in the south-western United States, some with species on the 
Mexican Plateau and the western Sierra Madre. One species of JJta [II. bicarinata) 
is found on the plateau and also on the west coast from Presidio to Tehuantepec. 
Sceloporus and Phrynosoma are important genera that range from the southern United 
States to Central America, and include species on the plateau as well as in the 
lowlands. 

The Geckonidse are found in all tropical countries ; in America they are almost 
entirely neotropical, and are not found on the Mexican plateau. The Geckonidae are 
peculiarly liable to accidental dis])ersal, and this is well illustrated by the distribution 
of Gehyra mutilata ; it is found on the islands of the Indian Ocean and the Malay 
Peninsula and Archipelago eastward to New Guinea, but it is also known from 
Western Mexico. The small family Eublepharidse, specialized Geckos, includes three 
species from Africa and India, distinguished by their stout form and by their enlarged 



EEPTILIA, BATEACHIA, AND PISCES. 115 

chin-shields from the four American species ; two of these, from Texas and California 
and from the West Coast of Mexico, differ in their scaling from the other more 
tropical species, one of which inhabits Panama and the other ranges from Costa Rica 
to Vera Cruz. 

The other families of Lizards found in America teach us nothing more than those 
dealt with above. There are several families common to the nearctic and neotropical 
regions, and most of these are found in other parts of the world also. South America 
is much richer in genera and species than North America ; most of the Lizards of 
Central America north to the outer slopes of mountains that border the Mexican 
plateau belong to or are related to South American genera, and most of those found 
on the plateau are generically distinct from them, but in some cases (e. g. Sceloporus, 
Phrynosoma) only specifically so. 

There is not, on the whole, a very fundamental difference between the Lizard faunae 
of North and South America; considering the differences in climate, and other 
features, we should expect most of the genera to be distinct, and in southern Mexico 
the transition from an elevated, dry, and almost treeless plateau bordered by mountains 
to a lowland district with tropical rainfall and vegetation accentuates this distinctness. 
But there are few differences that can be assigned definitely to the former isolation of 
the two continents, as practically nothing is known of the Tertiary history of the 
group. We know that some Mammalian genera, now tropical, formerly ranged 
further north and migrated from Asia^ to North America and thence to South America 
in late Tertiary times. Doubtless some Lizards, such as Mabuia, had a similar 
history ; but whether the Teiidfe, for example, were formerly nearctic, or whether 
they are for the first time spreading northwards, is a question difficult to answer. 

Snakes. — The burrowing snakes of the genus Typlilops have a wide distribution in 
the tropics ; there are a few species in Mexico and Central America, and one of these 
is T. brahminus, collected by Dr. Gadow in Michoacan. Previously this species was 
known to range from Africa to China and on islands eastward to the Moluccas; 
probably it will be found on some of the islands of the Pacific. This is an interesting 
parallel to the Gecko Gehyra mutUata ; in both cases Indian species have reached the 
West Coast of Mexico, and appear to have established themselves. The occurrence of 
Typlilops on oceanic islands indicates that the wide range of this genus is not due to 
its antiquity, but to susceptibility to accidental dispersal. 

Another genus of burrowing snakes, Glauconia, is found in America, Africa, and 
soutli-western Asia. G. alhifrons is tropical, ranging from Argentina to southern 
Mexico, but other species are found on the Mexican plateau and in the southern 
United States. 

The Boidae show some points of interest. The Pythoninae include a score of species 
from Africa, India, and Australia and a single American species, Loxocenus bicolor, 

<i2 



116 OKIGIN, ETC., OF THE FAUNA. 

from Guatemala and southern Mexico. The Boinse are widely distributed in the 
Old World, but the majority are American, and the occurrence in Madagascar of 
species of the Tropical American genera Corallus and Boa is curious ; it is evident 
that these genera had formerly a much wider range. 

Many other genera of snakes have a wide distribution ; thus the Crotalidse have 
several genera common to Asia and America, and, although they extend southwards 
to the La Plata, there are no peculiar Neotropical genera. One species of Crotalus 
ranges from the United States to Argentina. 

Without going into further details, it may be said that, as compared with the Lizards, 
a larger proportion of the Central American genera are found also on the Mexican 
plateau and in the southern United States, although there are, of course, a number of 
neotropical genera that range northwards only to southern Mexico. 

The Amblycephalidse are the only family of snakes that are strictly neotropical in 
America ; there are three South American genera, one of these, Leptognathus, having 
several Central American species, the northernmost in Tehuantepec. The other two 
genera occur in South-Eastern Asia, and it is probable that the family was formerly 
a northern one. Indeed, it is not unlikely that the whole neotropical Ophidian fauna 
has been derived from immigrants from the north that did not begin to reach South 
America until the Miocene connection was established. The distribution of the vast 
family Colubridse seems to favour this view. 

REPTILIA, BATRACHIA, and PISCES. 

Smnm^ry. — The mountains that fringe the Mexican Plateau form the boundary 
between the Neotropical and Nearctic Regions. True fresh-water fishes can neither 
cross seas nor travel overland from one river to another ; consequently their dispersal 
has been slow. The nearctic and neotropical fishes are quite difierent, and belong to 
different families ; the two faunae scarcely overlap ; in the Lerma System, on the 
southern part of the Mexican plateau, there are no neotropical fishes, the nearctic 
fishes are mostly generically distinct from those of the Rio Grande, and two endemic 
groups that are neither nearctic nor neotropical are the most important elements. 

For most Batrachians the sea is an impassable barrier, but they can migrate over- 
land and their dispersal may have been rapid. The nearctic and neotropical 
Batrachians are mostly distinct, but overlap to a considerable extent ; a few genera 
[Bufo, Ilyla) range throughout both regions. The rapidity of dispersal as compared 
with the fishes is exemplified by the holarctic group Urodela ; one genus, Spelerpes, 
has penetrated far into South America, whereas only one nearctic fish, the ancient 
Lepidosteus, has reached Panama. The spreading of Amhlystoma tigrinum of the 
United States southwards to the border of the Mexican plateau is in striking contrast to 
the peculiarity of the fish-fauna of the southern half of the plateau, where no species 



REPTILIA, BATEACHIA, AJiD PISCES. 117 

of fish occurs that is found in the United States. Crocodiles and Tortoises, like many 
families of Anura, are ancient and widely distributed. Snakes and Lizards are more 
modem, and their dispersal seems to have been recent and rapid ; in these groups 
there is no marked difference between the nearctic and neotropical regions ; no 
orders or families that show much regard for the boundary; the differences are 
for the most part generic or specific, and are mainly due to the sudden change 
from the dry plateau of the southern United States and Mexico to the tropical 
forests of Central America. 

Of the groups dealt with in this chapter, only the Fishes give clear indications as to 
the past, and from them we infer a Cretaceous connection between South America and 
Africa that may have persisted into the Eocene, and was succeeded by a long isolation 
of South America and in the Miocene by its connection with North America, which 
has been since the Eocene until quite recently more or less continuously connected 
with Asia across the Bering Sea. 



[ 118 ] 



AEACHNIDA*. 

By R. I. PococK, F.R.S. 

Introduction. 

Discussions on the geographical distribution of the Arachnida, especially of 
Scorpions, commonly open with a reference to the great antiquity of the group, and 
to the possibilities thus afforded it of achieving cosmopolitan dispersal. Dr. Hans 
Gadow, for instance f , dismisses the Scorpions in the following paragraph : — " This 
group is a good illustration of the effect of great antiquity. Scorpions already existed 
in the Silurian, and even some existing species date back to the Coal Measures! 
They have had every chance of spreading widely. A species of Tityus is preserved in 
Miocene amber of the Baltic, this genus is now restricted to southern South America. 
The group is cosmopolitan, limited only by cold, yet it is absent from New Zealand. 
They show scarcely any generic affinity between the Old World and the New, nor 
between South America and Australia. They have had sufficient time to develop 
along lines aloof from each other in these great land complexes." 

Much of this is untrue both in substance and in fact. None of the many Carboni- 
ferous genera can be referred with certainty even to existing families ; and the record 
of a species of " Tityus " from the European Miocene can only be regarded as evidence 
of the existence of the Buthida; in the Baltic area in mid-Tertiary times ; and, as 
regards the distribution of existing forms, if the views above expressed be accepted, 
there is nothing more to be said upon the subject. They are, however, inadmissible 
since they leave wholly unexplained the fact, singular though it be, that the present 
distribution of Scorpions does not attest the great antiquity of this order. If they 
were not known to be of Carboniferous age they might be judged, from the analogy 
supplied by Mammals, to date from late Mesozoic and early Tertiary times, for, as I 
have elsewhere pointed out;|;, if the surface of the world be regionally divided in 
accordance with the distribution of Scorpions, the resulting map will agree tolerably 
closely with the map based upon the distribution of Mammals, due allowance being 
made for the absence of Scorpions at the present time from all countries to the north, 
roughly speaking, of the 45th parallel of north latitude. 

* The Opiliones and the Acari have been omitted from this essay, the former because Mr. F. 0. P. Cam- 
bridge was not sufficiently acquainted with them to make his results altogether reliable, and the latter 
because of the imperfection of our knowledge of the Aoarine fauna of other parts of the world. 

t ' The Wanderings of Animals,' 1913. 

t ' Natural Science,' iv. pp. 353-364, 1894, and 1899, pp. 213-231. 



AEACHNIDA. 119 

A striking Mammalian feature in the distribution of Scorpions is their absence from 
New Zealand. Another is to be found in the character of the Australian forms, some 
of which are quite peculiar, while others show affinity with genera and species from 
southern Asia, like the Australian Rodent Mammals, and one, like Thylacinus, belongs 
to a South American family. Again, some of the South Asiatic genera do not pass 
Wallace's Line, and the small number of Mascarene forms are related to, but distinct 
from, those of tropical Africa and Asia, like the Mascarene Civets and Lemurs. 
Similarly, the tropical African and Asiatic species, although related, belong mostly to 
distinct genera. 

What is true of the Scorpions is true also, broadly speaking, of the two suborders of 
Pedipalpi, the Urotricha and the Amblypygi, which, like the Scorpions, existed in 
Carboniferous times in Europe and North America. The absence of both from New 
Zealand and Australia, and of the Urotricha from Africa also, are suggestive rather of 
comparatively recent origin than of high antiquity. 

The same cannot, however, be said of a great many of the Araneae (Spiders). The 
most ancient existing type, Lij)histius, apparently related beyond doubt to Carboni- 
ferous genera, is restricted to Indo-Malaysia, whereas many genera, like Lycosa, 
Aranea, Tetragnatlia, and othei's, not known to be ancient forms, are practically 
cosmopolitan in distribution. Moreover, some groups of spiders — perhaps Mesozoic or 
even Caenozoic, but not, so far as records tell us, Palaeozoic — attest by their restriction 
to the southern Continents the former existence of " Antarctica " much more forcibly 
than the orders of Arachnida known to have been in existence in the Carboniferous 
Period. Another instance of the restricted distribution of a Carboniferous order is 
supplied by the Rianulei, which are found now only in tropical West Africa and Brazil. 

Thus some of the Carboniferous Araclinida, the Scorpions, exist in Europe, Asia, 
Africa, America, and Australia; others, the Amblypygi, in Asia, Africa, and America; 
others, the Urotricha, in Asia and America ; others, the Rianulei, in Afnca and South 
America; the Mesothelid Spiders {Lijphisticus) only occur in the East Indies, while 
other spiders, assumed on morphological grounds to be of later date, may be cosmo- 
politan, and such spiders, when young, are known by their method of floating on 
webs to be able to cross arms of the sea in the direction of prevalent winds *. 

From these facts it seems clear that the present distribution of the Arachnida 
depends, not upon the duration of their existence, but upon their means of dispersal 
and power of adaptation to varied conditions. Hence the attempt to explain away 
the facts of their distribution by an appeal to their antiquity is barren of results, and, 
since the palaeontological history of the Arachnida throws no useful light upon the 
matter in hand, it has been ignored in the following discussion. 

• This means of dispersal is analogous to the flight of birds, whose powers to eross traets of ocean cannot 
be ignored in discussions of geographical distribution. 



120 OEIGIN, ETC., OF THE FAUNA. 

Order SCORPIONES *. 

The genera of this order recorded from Central America may be classified, with 
their distribution, as follows : — 

Family ScoRPioNiDiE. 

Subfamily Ischnurinae. 

Genus Opisthacanthus. Panama, Colombia, W. Indies. 
Subfamily Diplocentrinse. 

Genus Diplocentrus. Texas, Mexico, W. Indies, Brazil. 

Family VjEJOviOiE. 

Subfamily Vaejovinse. 

Genus Hadrurus. S. States of N. America ; Mexico, ? Guatemala. 

Genus Veejovis. S. States of N. America j Mexico. 

(Genus Uroctonus. California ; ? Guatemala.) 

(Genus Anuroctonus. California, Utah, etc. ; ? Guatemala.) 
Subfamily Megacorminae. 

Genus Megacormus. Mexico. 

Genus Plesiochaclas. Mexico. 
Subfamily Chactinae. 

Genus Broteochactas. Panama, Guiana. 

Family BuTHiDiE. 

Genus Centruroides. S. States of N. America, Central America, Antilles, S. America. 

(Genus Rhopalurus. Cuba, Haiti, Brazil, ? Mexico.) 

Genus Tityus. Mexico, Costa Rica, Panama, W. Indies, S. America. 

(Genus Isometrus. Commercially imported, if present.) 

With the substitution of the older name Veejovidae for luridae and the introduction 
of the MegacorminsB, containing genera unknown to me at the time, this is the 
classification proposed by myself in 1893 (Ann. Mag. Nat. Hist. (6) xii. pp. 305-311) 
and adopted in my paper on "The Geographical Distribution of Scorpions" in 1894 
(Nat. Science, v. pp. 353-364). In his contribution to the same subject published in 
1905 (Zool. Jahrb. Syst. xxii. pp. 321-364), Kraepelin suggested certain modifications, 
erecting the Diplocentrinse, Vaejovinae, and Chactinae to full family rank, including the 
Megacorminse as a subfamily of the Chactidae, and making Centruroides, with 
Rhopalurus, and Tityus, the types of separate subfamilies of the Buthidse. These 
amplifications, whether desirable or not, are of no importance in the present 
connection. 

Some of the genera above enumerated have been placed in brackets because 

• Although it is customary to designate the primary divisions of the Arachnida as " Orders," it must be 
remembered that the morphological differences between the groups of that rank are greater than those which 
distinguish the " Classes " of the Vertebrata. 



ARACHNID A. 121 

the evidence for tlieir occurrence in Central America is doubtful and requires 
confirmation *. 

Distribution of the Families and Subfamilies of 
Central American Scorpions. 

No genera of Scorpions are common to the Old and New Worlds, except hometms, 
one species of which has been commercially carried everywhere from the East Indies. 
None of tlie families mentioned above, however, are peculiar to America. The Old 
World appears to be the headquarters of the Scorpionidee, since this family is 
represented there by many genera distributed over Africa, South Asia, and Australia 
and referable to four subfamilies, only two of which, the Ischnuringe and Diplocentrinse, 
occur also in America. 

The Va?jovidae, absent from Africa and Australia, range from the Mediterranean 
area eastwards through North India into Further India, and reappear in North, 
Central, and South America, but are absent from the Antilles. Unless the Mediter- 
ranean genus Turns and the Indian genus Scorpiojps be included, as has been done, 
in the Vsejovina^, the American subfamilies are peculiar to the New World. But, in 
any case, the distinctions between the subfamilies of this family are finely drawn. 
Excluding the two Old World genera above mentioned, the Vaejovinse contain the four 
American forms enumerated above and two from South America. No other genera of 
the Megacorminae are known. The Chactinae, however, are represented by several 
genera in the northern countries of South America. 

The BiithidcTe are mainly an Old AVorld group. The family contains a large number 
of genera and species in Europe, Africa, Madagascar, South Asia, and Australia. 
Apart from the aberrant South American genus Ananteris, the only known American 
genera are the three indigenous forms enumerated above. These do not differ more 
from each other, or from some of the Old World genera, than the latter differ from 
each other. There is no evidence, indeed, that Tityus is not closely allied to the 
Oriental genus Isometrus. 

* The genera Uroctonus and Anuroctonus, for example, were included in my report on the testimony of 
Thorell, who recorded Uroctonus mordax and Anuroctonus jphceodactylus, two species originally described from 
California, from Guatemala. The specimens he had for examination, however, were sent to him with an 
example of Uadrurus Jiirsntus, also a Californian species, by Dr. Gustav Eisen, who was then living at 
San Francisco, and my opinion that they were wrongly localised is based upon the improbability of identical 
species inhabiting the two countries in question. The Buthid genus lihopalurus was included on the eddence 
of old specimens of lihopalurvs junceus, ticketed " Mexico " in the British Museum, the species in question 
having its home in Haiti. Finally, although Isometrus no doubt exists in the seaport towns of Central 
America, it has not yet been actually recorded, and as a commercially introduced form is wholly without 
interest. 

BIOL, centr.-amee., Introd. Vol., June 1915. B 



122 OEIGIN, ETC., OF THE FAUNA. 

General Features of the Scorpion Fauna of Central America. 

The nine genera of Scorpions certainly ascertained to be Central American are 
faunistically assignable to a northern category, a southern category, and a category 
which is botli northern and southern. 

To the southern category belong Opisthacanthus, occurring in Panama and Colombia ; 
Broteochactas, also found in Panama and the northern countries of South America 
(Guiana, etc.) ; and Tityus, which ranges all over South America from the Argentine 
northwards and is found in Panama, Costa Rica, and Mexico. The Mexican species, 
however, is very little known. It is not closely allied to the species from Costa Rica 
and Panama, which are identical with, or very nearly related to, species from the 
Amazons, Demerara, etc. Similarly, the species of Opisthacanthus and Broteochactas 
occurring in Panama are the same as species found in South America. 

To the northern category belong Iladrurus and Vcejovis, which extend from the 
Southern States of North America into Mexico, Vcejovis being one of the dominant 
Scorpions in the latter country. It is interesting to note the wide geographical 
severance between these two and their nearest allies, Carahoctonns and Iladniroidcs, 
which inhabit Ecuador, Bolivia, and Chile. Also to this northern category must be 
referred the two genera of Megacorminae, Megacormus and Plesiochactas, which are 
found in Mexico (Vera Cruz, Cordova), the latter extending to Guatemala. Their 
nearest allies are the Chactinse of tropical South America. 

To the category which is both northern and southern in distribution belong the 
genera Liplocentrus and Centruroides. The former ranges from Texas into Mexico, 
but not, so far as is known, to the south of the latter country in Central America. 
Nevertheless, it is found in the Greater and Lesser Antilles and in the northern part 
of South America (Brazil). Centruroides, on the other hand, occurs in the southern 
states of North America, throughout Central America, in South America as far down 
as Chile along the western side, in Brazil, and in the Greater and Lesser Antilles. 

Thus of the Central American genera only four occur in the Antilles, Tityus, 
Centruroides, Biplocentrus, and Opisth acanthus. 

Two of the above-mentioned genera, Opisthacanthus and Diplocentrus, are of 
outstanding interest from the geographical point of view, because of the singular 
distribution of the subfamilies to which they belong. 

Opisthacanthus is unmistakably related to the tropical and Soutli African genus 
Opisthocentrus. The generic distinction between the two is not, indeed, always 
admitted. Tropical Africa is also the home of several more genera of the subfamily 
Ischnurinse, while others occur in Madagascar and South Asia as far east as 
Australasia. The kinship between the African and the solitary monotypical tropical 
American genus suggests that South America received this element of its fauna by 
means of a direct land-connection with Africa. 



AEACHXIDA. 123 

The distribution of the Diplocentrinae is still more remarkable. Four genera belong 
to this subfamily, two being American and two Arabian. In addition to Biplocentrus*, 
the range of which has already been stated, America contains the well-marked genus 
Oiclus, known hitherto only from Antigua in the Lesser Antilles. The Arabian genera 
are Neho recorded from Syria and Arabia and Ileteroncbo discovered in the island 
of Abd-el-Kuri by Sokotra. This singular discontinuity in distribution is closely 
paralleled by that of the Centipede Scoloj^endra (see p. 134). To explain it one may 
tentatively suggest a transatlantic connection between the Mediterranean area of 
Europe and America by way of the Antilles, an hypothesis supported by the survival 
of Scolopendra in the Canary Islands. Assuming this to be the explanation, it is 
probable that America received its Diplocentrine Scorpions from the Mediterranean 
area, and not vice versa, because the Old World, and not the New World, is at the 
present time the home of the Scorpionidise, to which the Diplocentrinae belong. 

Very little can be said about the origin of the remaining Central American genera. 
The absence of the Vajjovidse from tropical Africa eliminates the latter country as a 
possible source for this family. Its mainly extratropical range in the Old World, 
coupled with its existence at the present time in North America, suggests its former 
wide distribution in the Northern Hemisphere ; and possibly the break in the 
distribution of the VsejoviniE in America, above alluded to, points to the southward 
migration of the family from the western parts of North America to the western parts 
of South America before the elevation of the isthmus of Panama. In this connection 
the absence of the family from the Antilles is significant. 

The dominance of the Buthidfe in the Old World as compared with the New 
World suggests the Old World as the original home of the family ; but whether the 
ancestors of the American genera entered America by a transatlantic bridge from 
Africa or Europe or by the known eastern or western northern routes under tropical 
or warm temperate conditions, it is impossible to say. 

Rkgional and Subkegional Distribution of American Scorpions. 
Making allowances for incomplete knowledge, which renders accurate mapping of 
boundaries impossible, it seems that America may be divided by its Scorpions into the 
same geographical areas as those indicated by the Mammalia. The northern parts 
of the continent to the north, roughly speaking, of the 40th parallel agree with the 
corresponding areas of the Old World in being without Scorpions. This negative 
feature defines the Holarctic Region. Sharply distinguished from this by the 
presence of Scorpions are the Southern States of the Union ; and the extension of two 
of the genera {Hadrurus, Vajovis) into Mexico, but no farther, affiliates the greater part 

• Including Didymocentras, Kraepelin. 

e2 



124 OEIGIN, ETC., OF THE FAUNA. 

of that country with the southern United States to constitute the Sonoran Region, 
which corresponds with the Mediterranean Region of the Old World. Two more 
genera, XJroctonus and Anuroctonus, are peculiar to the Sonoran, but have not been 
recorded from Mexico. Probably also Plesiochactas and Megacormus from Mexico and 
Guatemala belong to it. The other Sonoran genera, Diplocentrus and Ceiitruroides, 
extend far to the south, and link the Sonoran Region with the Antilles and not 
South America. 

South America, the Neotropical Region, is characterised by the family Bothriurida;, 
ranging from Peru, Chile, and Brazil to Patagonia ; by a large number of genera and 
species belonging to the Chactinae, mainly restricted to the northern parts of the 
continent, one genus Broteochactas spreading into Panama; by the Ananterinae, a 
peculiar subfamily of Buthidse ; by the Buthoid genus Tityus, which spreads through 
Panama and Costa Rica to Mexico ; by the peculiar genus of IschnurintE, OpistJtacanthiis, 
which occurs in Colombia and Panama ; and by two peculiar genera of VsejoviuBe, 
Carahoctonus and Iladruroides, ranging from Chile to Ecuador. Thus the southern 
portion of Central America contains Scorpions generically, in some cases specifically, 
identical with those of the northern countries of South America. If it be separated 
as a subregion from South America, an immaterial point, it may be defined by the 
■absence of a large number of South American genera. 

The affinities of the Antilles are certainly with Neotropical rather than with the 
Sonoran Region. Two Sonoran genera, Diplocentrus and Cerdruroides, it is true, occur 
both in the Greater and Lesser Antilles, but both are found in South America as well. 
The genus Ehopaltirus, allied to Centruroides, and occurring in Haiti and Cuba, is 
also found in Venezuela, Brazil, and Colombia; Opisthacanthus, inhabiting Colombia 
and Panama, has been recorded from Haiti ; and the essentially Neotropical genus 
Tityus exists both in the Greater and Lesser Antilles. One genus, namely Oiclus, is 
peculiar to the Lesser Antilles *. Judged from their Scorpion fauna, therefore, the 
Antilles may be regarded as a subregion of the Neotropical Region, characterised 
mainly by the absence of some conspicuous South American types, e.g. the Chactinae 
and the Bothriuridae. 

Evidence supplied by the Scoepions for a Land-connection between 
South America and Australia. 
Although the following fact has no direct connection with the Scorpion fauna of 
Central America, it is sufficiently interesting to call for mention, since it strongly 

• In my paper on the Scorpions of the West Indies (Journ. Linn. Soc, Zool. xxiv. pp. 374-404, 1893), 
the genera lladrwus and BrachistosUrnus were recorded from these islands. The specimens of Iladrurus 
and Brachistcsternus were certainly wrongly labelled. The former, in the lierlin Museum, is, as Kracpelin 
has shown, an Hadruroides from Ecuador ; and the specimen of Bracldsiosternus, one of the Bothriuridae, is 
identical with a well-known Peruvian species. 



AEACIINIDA. 125 

confirms evidence supplied by other groups for a direct land-connection by way of the 
Antarctic or the South Pacific between South America and Australia. 

The family of the Bothriuridae is represented in South America by many genera and 
species, occurring mainly in Patagonia, Chili, the Argentine, Buenos Ayres, and Brazil. 
Beyond the limifs of South America the family is unknown, save for the occurrence 
of the single genus and species, Cercophonius squama, which is found all over 
Australia, apart from its northern portions, and in Tasmania. Since South America 
is obviously the headquarters of the family, it may be inferred that this element of 
the fauna of Australia was received directly from South America ; but since this 
Scorpion, like all Scorpions, is absent from New Zealand, it may be further inferred 
that New Zealand formed no part of the laud connecting Australia and South 
America at the time the ancestor of the Scorpion in question migrated from 
South America. 

The Scorpions thus supply evidence for a transatlantic connection between tropical 
Africa and South America and for a transpacific connection between South America 
and Australia : by means of the former America may have received certain elements 
of its Scorpion fauna, by means of the latter it may have contributed something to 
the fauna of Australia. 

Order PEDIPALPI. 

The existing Pedipalpi of the suborder Urotricha are referred to a single family, 
the Thelyphonida;, which extends in the Old World, where it is represented by many 
genera, from India to Austro-Malaysia and northwards to China and the Philippine 
Islands ; and in America from the Southern States of the Union to Brazil. Two 
genera only are known from America, Thelyphonellus from the Amazons and Mastigo- 
procUis. The latter is represented by a few species in Brazil, a few in the Lesser and 
Greater Antilles and two in Central America, one recorded from Guatemala, the other 
from Mexico (Cordova, Jalisco, Guerrero, etc.). The Mexican species extends into 
Texas and adjoining States of North America. 

The nearest ally of Mastigoproctus appears to be the eastern Himalayan genus 
Uroproctus. This fact, coupled with the occurrence of the family in China, points to 
its former existence throughout the countries bordering the North Pacific, when tropical 
or warm temperate conditions prevailed there. If this be so, the tropical American 
forms must be regarded as immigrants from the north. 

One point connected with the distribution of the group in Central America is its 
apparent absence from the States to the south of Guatemala. Nevertheless, it occurs 
in the Greater and Lesser Antilles (Haiti, Martinique) and in Brazil. Possibly 
therefore it made its way from Central into South America by means of an Antillean 
route. 



126 OllIGIN, ETC., or THE EAUNA. 

The Amblypygoiis Pedipalpi have a wider range than the Urotricha. In the Old 
World they are represented by two families, the Tarantulidse ( = Phrynichid8e) which 
range from Siam, through India, and Arabia into tropical and South Africa, but are 
absent from Madagascar, and the Charontidae which have their headquarters in 
Further India, whence they spread westwards to the Seychelles and eastward to the 
Solomon Islands. Like the Urotricha they are absent from Australia *. 

The suborder is represented in tropical and subtropical America by tlie family 
Phrynidee, which is unknown in the Old World. Of the two subfamilies, the 
Heterophryninae are essentially South American, but are unknown in the Antilles. 
One species, however, which for want of accurate determination I have provisionally 
referred to the DemeJraran //. chiracanthus, has been recorded from Central America 
without further particulars as to locality. 

The Phrynina? have a much wider range, extending from North America, throughout 
Central America and the West Indies, into South America. Acanthojjhri/nus is known 
only from California and Mexico (Durango). Of Phrynus, one species ranges from 
Texas to Nicaragua ; two occur in Guatemala and are not known elsewhere, and one in 
Panama, the latter being found also in Colombia, Venezuela, Trinidad, and Barbados. 
Other species occur in the Lesser and Greater Antilles and the northern part of South 
America. The third genus, Ilemiphrynus, ranges from Mexico (Tuxtla, Oaxaca, Teapa, 
etc.), through Guatemala, Costa Rica, and Panama into South America. Although 
unknown in the Antilles, there is an outlying species in the Bahamas. 



Conclusion. 

The Pedipalpi supply no evidence of a mid-Atlantic land-connection between Africa 
and America, and none of the existence of a continuous or partial Antarctic land 
joining the southern continents. Nor in the case of America do they help much in the 
definition of the regions and subregions indicated by the Scorpions. One or two points 
may, however, be touched upon. 

The occurrences of Acanthophrynus in California and Mexico, of Phrynus vjhitei in 
Texas and Mexico, of Mastigoproctus giganteusf in several of the Southern States of 
America and all over Mexico, affiliate the latter country faunistically with the southern 
portions of North America. The Sonoran area thus defined difl^ers from South 

* One species of Urotricha belonging to the genus Thelyphonus has, however, been recorded from Cape 
York, the extreme northern point of Australia. 

t Since the Pedipalpi and Scorpions have existed from the Carboniferous with comparatively slight 
structural changes, they cannot be regarded as plastic organisms. In my opinion, therefore, species, genera, 
and families have greater significance from the geographical standpoint than groups of those ranks in tlie 
Mammalia. 



AEACFNIDA. 127 

America in the absence of Ileterophrijnus and of the Thelyphonid Thehjphonellus ; but 
it is linked thereto by the extension of Hemiphrynus from Mexico to Colombia, and of 
Plirynus and Mastigoproctus from Texas at least as far as Brazil. The chief distinction 
between the Antilles and the Sonoran area is the absence of Acanthophrynus from 
those islands, and between the Antilles and South America the absence oi Hetero- 
plinjnus, Thelyphonelhis, and II emi plirynus from the former. But the occurrence of 
Phrynus and Mastigoproctus in both the Greater and Lesser Antilles, as well as in 
Mexico and South America, points to union between these islands and the countries 
in question. 

Order SOLIFUG.E. 

With one exception the Central American genera of this Order show the same 
general geographical features as the Pedipalpi and some of the Scorpions. 
Eremohates, the type of a special subfamily, is restricted to the Southern States of 
North America and Mexico, its distribution coinciding tolerably closely with that of 
the Scorpion Vcejovis in the sense that neither extends into the West Indies or into 
South America. Ammofrecha, on the other hand, which also occurs in the Southern 
States of the Union, is found in the Antilles and in South America, its distribution 
being parallel to that of the Pedipalp genera Mastigoproctus and Phrynus and of 
the Scorpion Centruroides. The exception above referred to is Ilemiblossia, one 
species of which occurs in Guatemala and the others in South Africa. This case of 
discontinuous distribution recalls that of the Scorpion Opisthacanthus, whose nearest 
allies are tropical and South African species. 

Ammotrecha and Ilemiblossia belong to a subfamily represented by numerous genera 
in Africa, South-western Asia, and the Mediterranean area of the Old World, but 
unrepresented in the Oriental Region and Eastern Asia. Their occurrence in Central 
America points to an Atlantic connection between the Old and New Worlds. The only 
other facts to be borne in mind in this connection are the records of a species of Dcesia, 
typically an African and Mediterranean genus, in Mexico, and of a species of the 
genus Eremobates in Afghanistan. Both these records, however, require confirmation. 



Order ARANE^. 

The Central American fauna of Spiders differs, as a whole, from that of the Scorpions 
and Pedipalpi in two particulars, namely, the vast number of species, genera, and 
families it is composed of and the large number of these that are common to the Old 
and New Worlds. Hence, within the limits of a short essay, it is impossible to deal 
with all the families concerned. To illustrate some of the main features of the fauna 
and to trace as far as may be its sources, I have selected therefore for detailed treatment 



128 OEIGIN, ETC., OF THE FAUNA. 

the sharply circumscribed suborder of the Mygalomorphse. The mutual affinities of 
the genera and families of this group are sufficiently well understood for the purpose, 
and from the geographical standpoint it has the additional advantage of being, so far 
as wo know, dependent upon continuous land-areas for its dispersal, since the species 
appear — for the most part, at all events — to be independent of the method of travelling 
commonly known as " ballooning," which the young of many of the families of 
Arachnomorphae have been ascertained to practise shortly after dispersing from the 
cocoon. As has been suggested above (p. 119), the very wide, sometimes cosmopolitan 
distribution of some genera may perhaps be assigned to this cause, coupled with 
exceptional power of adaptation to highly varied conditions. 

The Central American representatives of the Mygalomorphse were referred by 
Mr. F. Cambridge to the three families, Clenizidae, Dipluridse, and Theraphosidte. 
This classification will be here adhered to in the main as sufficiently exact for the 
purpose in hand. 

The first subfamily of the Ctenizidae, the Actinopodinae, is represented by the two 
genera, Actinopus and Neoctewza. The latter has only been recorded from Guatemala 
and Demerara. The former is abundant all over South America from the Argentine 
northwards to Venezuela, and enters Central America at Panama. Perhaps it awaits 
discovery in other tropical districts of Central America. It is interesting to record that 
the only other representative of this subfamily is the Australian genus Missulena 
(Eriodon). 

The genera of the second subfamily, the Ctenizinae, are referred to three sections, 
the Pachylomereae, the Ctenizeae, and the Cyrtauchenieae. From the first of these 
should certainly be dismembered the genus Ch.orizo]}s as the type of a very special 
group, the Cyclocosmeae or Halonoprocteas. This group contains the three genera: 
Chorizops recorded from Mexico (Guanajuato, Vera Cruz), Ct/clocosmia from Alabama, 
and Jlalonoproctus from China. 

The Pachylomereae comprises three genera : Conothele ranging from Burma to the 
Solomon Islands, Ilehestatis from California, and Pachylomerus itself. This genus has 
a remarkably discontinuous distribution. Several species occur in the Southern States 
of North America, in Central America from Mexico southwards to Guatemala and 
Costa Pica, and a few in the West Indies (Jamaica, St. Vincent). In the Old World 
it is met with in Japan, but, singularly enough, elsewhere only in the western 
Mediterranean (Spain, Algeria). 

The genus Bothriocyrtum, the sole Central American representative of the Ctenizeae, 
has one Mexican and one Californian species. Its nearest ally is Cyrtocarenum of the 
Mediterranean area. All the other genera of the Ctenizeae are Old World forms, two, 
in addition to Cyrtocarenum, being Mediterranean, one South African, one Central 
Asian, and one Japanese and Chinese. 



AEACHNIDA. 129 

The Cyrtaucheniese, the fourth and last group of the Ctenizinte, has three Centrad 
American representatives, Eutychides, Eucteniza, and Enrico, all restricted to Mexico. 
One other genus, MyrmekiaphiJa, from Carolina and Virginia, occurs in the New World. 
Of the rest, four in number, one is Burmese, one (Cyrtauchenius) Mediterranean, and 
two South African. 

The Central American forms of the Diplurida? are referred to two subfamilies, the 
Diplurinffi and the Ischnothelinse. The Diplurinse, corresponding with Simon's group 
Diplureae, have a very singular distribution. In South America there are about nine 
genera ranging from Chili northwards, Fufiiis the single Central American genus 
spreading into Costa Rica and Guatemala from Brazil, Ecuador, and Bolivia. Four 
genera occur in Australia, two in Madagascar, and one in the Mediterranean area. 

The distribution of the Ischnothelinae ( = Macrothele8e, Simon) is equally remarkable. 
There are two known Central American genera. Evagrus spreads southwards from 
Mexico and Guatemala to Bogota, and northwards into the United States (Idaho). 
One species, possibly generically distinct from it, has been recorded from South Africa. 
The nearest ally of Evagrus appears to be the Transcaspian genus Phyxioschcema, which 
is itself closely akin to Stenygrocercus from Australia and New Caledonia. Other less 
closely related genera occur in the Old World in the Mediterranean area, in tropical 
West Africa, Burma, China, and New Zealand. Ischnothele is a very distinct type 
from those above mentioned. It occurs in Mexico, Nicaragua, Guatemala, and the 
northern countries of South America, and in the Old World, in tropical Africa, 
Madagascar, and India. 

The Central American genera of Theraphosidae * have been revised since 
Mr. Cambridge described them. The species he referred to Eitrypelma have been 
split up into a number of genera and are probably susceptible of further subdivision f . 
The species named cemilia, ■pallidum, smithii, and vagans belong to Brachypelma, 
which ranges from Mexico to Colombia in South America. 'I'hose named rusticum 
and seemani are referred to Aphonopelraa, Avhich extends from the Southern States of 
North America into Mexico and Costa llica. The species described as Eurypelma 
longipes from Guatemala is the only known form of the genus Cithar acanthus, which 
is allied to Cyrtopholis, a genus mainly confined to the Greater and Lesser Antilles, 
but represented in Central America by C. pernix, referred by Cambridge to Ilapalopus, 
from Orizaba in Mexico. The genus Dugesiella, containing species from Texas and 
Mexico (Guanajuato) has been added to the Central American fauna since 

• The Spiders of this family are those commonly referred to in textbooks and Natural Histories as 
Mygale, or " Eird-eating " Spiders. 

t One of the species, spntxdatam, was included owing to an error in locality. It belongs to the genus 
Orammostola (= Citharosceliis) and came from CLili. Another Chilian species that was entered and must 
also be eliminated is Paraphysa manicata. 

BIOL. UENTR.-AMEB., Introd. Vol., June 1915. s 



130 OEIGIN, ETC., OF THE FAUNA. 

Mr. Cambridge finished his account of the Spiders. So, too, has the genus Psalmopoeus, 
which contains the Costa Rican species reduncus and others from Ecuador, Colombia, 
and Trinidad. 

The genera Sphcerohothria from Costa Rica, Sericopelma from Panama, and 
Schizopelma from Mexico and Costa Rica have not, so fai-, been discovered beyond 
the limits of Central America, but, like the others cited above, and the remainder 
recorded by Mr. Cambridge, they are related to South American or West Indian 
genera of this great family, and with one or two exceptions have no near kinship with 
the genera characteristic of tropical Africa, Asia, and Australia, which belong to distinct 
groups or subfamilies. Two apparent exceptions to this, however, must be mentioned. 
The South American genus Avicularia which extends into Panama belcmgs to a group 
to which also belongs the tropical West African genera Scodra and Heteroscodra ; and 
the genus Hemirrhagus, represented by one species, cervinus, in Mexico, seems to have 
its nearest ally in Cratorrhagus of the Mediterranean area. 
The conclusion suggested by the above-stated facts is this : — 

The Central American Mygalomorphse belong to two categories : (1) Those referable 
to groups which are in the main northern in distribution ; (2) those referable to groups 
which are in the main southern in distribution. 

1. To the northern group belong Pachylomerus, which occurs elsewhere in the 
West Indies, North America, Japan, and Spain ; Chorizops^ whose only known allies 
are found in North America and China ; Bothriocijrtum, which occurs also in North 
America and is related to genera from Japan, China, Central Asia, and the Mediter- 
ranean area ; Enrico, Eufychides, Eucteniza, akin to genera from North America, 
Burma, and the Mediterranean ; Evar/rus, ranging from North America to Bo"-ota, 
is allied to genera from the Mediterranean region, AVest Africa, Transcaspia, China, 
and Burma. In the New World none of these groups is found south of the equator. 
In the Old World, however, genera akin to Bothriocyrtum, Enrico, Eutychides, 
Eucteniza, and Evagrus occur in South Africa, and others allied to Evagrus in 
Australia and New Zealand. Perhaps in the future these may be geographically 
linked with the northern types, but so far as our knowledge goes they are isolated. 

2. To the southern groups belong Neocteniza, from Demerara and Guatemala, and 
its ally Actinopus, which ranges from the Argentine into the West Indies and Panama 
and is related to the Australian genus Missulena, but to no other genus of the 
Old World ; Fujius, extending from Brazil to Costa Rica and Guatemala, is related 
to many genera confined to South America and to others occurring in Madagascar and 
Australia, with one outlying form in the Mediterranean area ; Ischnofhele, ranging 
from the northern countries of South America into Mexico, is also met with in 
tropical Africa, Madagascar, and India; all the Central American genera of Thera- 
phosidae, some of which pass into the Southern States of North America, have their 
nearest allies in South America, where they occur at least as far south as the 



AEACHNIDA. 131 

Argentine and Chili. In the Old World their nearest allies, few in number, are 
found in West Africa and the Mediterranean. 

Tentative explanations of these facts may be illustrated by one or two cases. If the 
groups to which Chorizops and Pachylomerus belong are North American in origin, 
they may have extended thence southwards into Central America and northwards into 
eastern Asia. Pachylomerus, in addition, may have passed eastwards from America to 
Spain and Algeria, or its apparent restriction in the Old World at the present time 
to such widely sundered areas as the south-western Mediterranean and eastern Asia 
may be attributed to its former extension over Eurasia and its survival only in those 
two localities. Against this hypothesis can only be urged the impossibility of assigning 
any cause for its extinction in other parts of the Asiatic district. 

If the group to which Actmojms and Eucteniza belong was evolved in South 
America, these genera may have moved northwards into Central America ; and the 
restriction of an allied genus to Australia may be assigned to the entry of the group 
into that country from South America. 

Since tropical America is the headquarters of Ischnothele, judging from the number 
of indigenous species, it may be provisionally regarded as the original home of the 
genus. Thence it may have passed by a transatlantic bridge to Africa, and so to 
Madagascar and India. 

These four genera of Mygalomorphse, apart from others which might have been 
cited, supply evidence, such as it is, for direct land-continuity, under tropical or 
warm temperate conditions, between (1) North America and eastern Asia; (2) North, 
Central, or South America and the Mediterranean ; (3) South America and tropical 
Africa; (4) South America and Australia. 

Corroborative evidence for "Antarctica" is supplied by another family (or sub- 
family) of Mygalomorphse, namely the Migidse, which is related to the Ctenizidge, 
but does not occur in Central America. It contains eight genera, distributed as 
follows : — Moggridgea and Pcecilomigas, South and South-east Africa ; Myrtale, 
Micromesomma, and Thyroponis, Madagascar; Heferomigas, Tasmania; Migas, New 
Zealand ; and Calathotarsus, Chili. 

It appears to me to be impossible to assign any reason for the extermination of this 
group of Spiders in the northern portions of the continents to which they are now 
restricted, if the genera in question are the descendants of a family evolved sufficiently 
far to the north to have passed from the Old to the New World, or vice versd, along a 
North Atlantic or North Pacific route. 

Turning to the Aracliuomorphoe, one or two of the Central American families may 
be selected to illustrate the restricted range of some and the wide ransje of others. 

The Senoculidse and Acanthoctenidae, each represented by a single genus, are 
peculiar to America, ranging from South America northwards to Mexico, The 
Psechridas are represented by two genera, Psechrus and Fecenia, in the southern and 

s2 



132 OEIGIN, ETC., OF THE FAUNA. 

south-eastern parts of Asia, an area to which they were believed to be restricted until 
the discovery of the genus Me.tafecenia in Central America (Guerrero in Mexico). 
The Deinopidse (Deinopis), occurring in North, Central, and South America, is also 
found in tropical Africa, Madagascar, and South Asia. Of greater interest, because of 
their restricted range, are the families Sicariidge and Caponiidse, both very sharply 
defined groups, the latter being also in some respects highly specialized or progressive. 
The Sicariidae range in America from Chile up the western portions of the continent 
as far as Costa Rica, and are onlv found elsewhere in the world in South Africa. 
Similarly with the Caponiidge. Of the three genera, Nops occurs in South America 
and the Antilles, Caponina in the Antilles and Central America (Guatemala), while 
the related genus Caponia is South African. These families corroborate the evidence 
derived from other sources for a direct connection between Africa and South America. 

The most prominent Spiders in the Central American fauna are those belonging to 
the web-spinning species of the family Argyopidse. The distribution of some of the 
dominant genera of this family is as follows : — Nephila ranges from the warmer States 
of North America through Central America into South America, and is abundant all 
over tropical Africa, Asia, and Australia ; Gasteracantha, a hard-shelled spine-bearing 
genus, has nearly the same distribution as Nephila. The other hard-shelled spiny 
genus, Micrathena, is, however, restricted to America, where its range closely coincides 
with that of the two just mentioned, Argyope agrees closely with Nephila, but has 
a more northern distribution, at all events, in the Old World, where it reaches the 
Mediterranean area. Finally, the genus Aranea {= Epeira) is practically cosmo- 
politan, extending from north temperate, if not subarctic, latitudes southwards all 
over the continents. 

These genera illustrate, as clearly as others which might have been selected, the 
wide distribution of a large number of the genera of Arachnomorphse recorded from 
Central America. 



CHILOPODA, 133 

CHILOPODA, DIPLOPODA, AND PROTOTEACHEATA. 

By R. I. PococK, F.R.S. 

The Chilopoda and Diplopoda were in existence in the Carboniferous Period in 
Europe and North America. But since these ancient forms were different from their 
modern representatives, and since they afford no help to an understanding of the 
present distribution of these classes, it is needless to discuss them further, and useless 
to repeat what has already been said on this head in the introduction to the essay on 
the Arachnid a. 

Class CHILOPODA (Centipedes). 

From the geographical standpoint the Chilopoda or Centipedes are remarkable for 
the wide distribution of the orders and of most of the families into which the class is 
divisible. It is probable, however, that when the group has been studied in greater 
detail, the number of families, especially of the Geophilomorpha, will be considerably 
increased and their distribution better understood. The recorded Central American 
genera of Geophilomorpha and of Scutigeromorpha call for no special comment in the 
present connection, since they exhibit no special features throwing light upon the 
origin of the fauna of that country. The Lithobioraorpha and Scolopendromorpha are 
more interesting. 

The Central American species of the former order belong to the family Lithobiidae, 
which, although of wide distribution with a few outlying species of the genus Lithobius 
in the Southern Hemisphere (India, Australia), is mainly northern, being spread all 
over Europe, Central Asia, and North America, and represented by a vast number of 
species in these continents. Hence it seems that the Central American representatives 
spreading through Mexico to Guatemala must be regarded as a northern element in 
the fauna of our area. 

The classification of the Scolopendromorpha proposed in my report may be revised 
as follows : — 

Family Cryptopid^*. 

Subfamily Cryptopinae (Crypfopsf, Theatops, Otocryptops, Scolopocryptops), 

Subfamily Newportiinse {Newportia, Scolopendrides). 
Family ScolopendkidjB (Cupipesf, Rhombocephalus, Scolopendra, Parotostigmus, RhysidaX). 

* Brolemann's examination of the type of Scohpendropsis, showing that the genua is closel}' related to 
Pithopus, proves that the presence of 23 pairs of legs cannot be regarded as a character upon which a group 
of family rank can be based. I therefore follow Kraepelin in associating Otocryptops and Scolopocryptopi 
with Cryptops. 

t Added by Kraepelin to the Central-American fauna since my report was written. 

t In Kraepelin's Monograph of the Scolopendridce (Jahrb. Hamb. Wissen. Arcb. xx. 1903, p. 139), this 
genus, as expanded by that author, should have been named Trematoptychus, -which antedates Rhysida by 
one year. 



134 OEfGIN, ETC., OF THE FAUNA. 

Of the family Cryptopidee the genus Cryptops is cosmopolitan, and its discovery in 
Central America has no particular significance. Theatops, Otocryptops, and Scolopo- 
cryftops are more restricted in range. Theatops spreads from North America into 
Mexico, and has one species in the Mediterranean area. Both Otocryptops and 
Scolopocryptops are South, Central, and North American, Antillean, and Chinese, 
Otocryptops also extending in the Old World into the south-eastern portion of the 
Oriental Region (Philippines, Celebes, Java, New Guinea, etc.) *. 

The Newportiinae are peculiar to the New World, being plentiful in South and 
Central America and the Antilles and represented by one species in North America. 
The two genera referred to the subfamily appear to be specialised forms of Crypto- 
pidse; and, judging from their present distribution, they were probably evolved in 
tropical America. 

Two of the genera of Scolopendridae demand particular attention — namely, Paroto- 
stigmus and Scolopendra, The former is mainly tropical American, occurring in 
South and Central America and the Antilles, but a few allied forms inhabit tropical 
Africa. Of greater interest are the species to which in 1903 (Nat. Hist. Sokotra, 
p. 429) I restricted the term Scolopendra +. This genus, as here limited, ranges from 
the Southern States of North America, through Central America and the Antilles, into 
the northern parts of South America and the Galapagos Islands. Elsewhere it has 
only been recorded from the Canary Islands, the Kameruns, Arabia, Syria, Persia, 
Sokotra, and the adjoining island of Abd-el-Kuri. Its distribution, therefore, coincides 
very closely with that of the Diplocentrine Scorpions, and is suggestive of a direct 
land-connection across the tropical Atlantic between the Old and New Worlds. 

The Chilopod fauna of Central America shows marked affinities with that of North 
America, Asia, and Europe (LUhobius) ; with Eastern Asia {Otocryptops, Scolopo- 
cryptops) ; with tropical Africa {Parotostigmus) ; and with the Mediterranean area 
{Scolopendra, Theatops). But there are no data justifying any conclusion as to the 
original home of the genera in question, unless superiority in number of species be 
regarded as a criterion on that head. If this somewhat unsafe basis for an opinion 
be adopted, the inference is that all the genera, except the northern LUhobius, were 
evolved in tropical America, and subsequently entered the continents of the Old World 
where they are now found. 

• One species of Otoci-i/ptops, abundant in South and Central America and the Antilles, has also been 
recorded from West Africa. There is no proof, however, that the species was not transported, probably in 
connection with the slave trade, as I have suggested. 

t Nearly all the Central American species belong to this genus. The two aberrant torviiB, punctiventris and 
pygmaa, should perhaps be regarded as representing distinct genera and need not be considered in the present 
connection. 



DIPLOPODA. 135 



Class DIPLOPODA (Millipedes). 

The dominant Central American forms of the Calobognatha or suctorial millipedes 
belong to the genus Platydesmus of the family Platydesmidse. This family also has 
representatives in North America, Amurland, Malaysia, and the Mediterranean area 
in Europe. It has not been recorded as yet from tropical Africa, India, or Australia. 
Hence the known facts of its distribution point to its evolution in the Northern 
Hemisphere and to its failure to penetrate into the ancient southern continents. 
If the Mediterranean species referred to Platydesmus belong, as is probable, to a 
distinct genus, Platydesmus is restricted to Central America (Mexico, Guatemala). 

The Chordenmoidea are divisible into two families — the Heterochordeumidae, 
ranging from India to New Zealand, and the Chordeumidae, which are abundant in 
North America and Europe. Their range eastward from Europe into Asia has not 
been ascertained, but there seems no doubt that they are absent from tropical Africa. 

The described Central American species, from Mexico and Guatemala, belong to a 
genus (Cleidogona) also existing in North America. No species have been discovered 
in South America or in the Antilles. This element, therefore, of the Diplopod fauna 
of Central America was no doubt derived from North America. 

The Stemmiuloidea are an obscure group, known at present only from tropical Asia 
(Ceylon), tropical Africa (Liberia), and tropical America (Colombia, Panama, and 
Porto Rico). Probably these Diplopods await discovery elsewhere ; but, so far as is at 
present known, they may be described as southern. Perhaps the most interestin;; 
point connected with their distribution is the ascription to the same genus (Diopsiulus) 
of the Ceylonese, Liberian, and Porto Rican species. This classification accords 
with the theory of the former existence of a tropical transatlantic connection between 
the Greater Antilles and West Africa. 

The Central American forms of luloidea, Paraiulus, from Mexico and Guatemala, 
are closely akin to genera and species now living in North America. The nearest ally 
of Paraiulus in the Old World is Mongoliulus, recorded from Corea. Here again 
there is evidence for a northern derivation of this element of the Central-American 
fauna. 

As an explanation of the occurrence of the suborder in North America and Europe, 
it may be suggested that it formerly extended across the Northern Atlantic when 
temperate conditions prevailed in Greenland, Iceland, and the now-vanished land-areas 
which are believed to have joined these countries and Europe and America in one 
continuous tract. 

Of the Spirostreptoidea, the single Central American genus of Spirostreptidae 
{Orthoporus) is mainly South American in distribution; but it is closely allied to, if 
not identical with, millipedes of the same family now found in tropical Africa, but not 



136 OEIGIN, ETC., OF THE FAUNA. 

apparently in tropical Asia. The most plausible explanation of these facts is that 
Orthoporus and its near allies were evolved in a continent embracing and connecting 
South America and tropical Africa, and that Orthoporus moved northwards into 
Central America after the formation of the isthmus of Panama. In Central America 
the genus ranges from Panama to Mexico. 

Like Orthojwrus, the genus Epinannolene also appears to be South American in its 
affinities. 

Although the precise geographical range of the two dominant Central American 
genera of Spiroboloidea, namely Bhinocricus and Spiroholus, has yet to be ascertained, 
one or two facts of interest may be noticed. 

Spirobolus, ranging from Mexico to Guatemala, occurs also in the Southern States 
of North America and in Eastern Asia as far north as Pekin. It is not abundant in 
the West Indies, although recorded from Porto Rico. It is apparently absent from 
South America. The evidence, therefore, is in favour of a northern, rather than a 
southern, origin for this genus. 

Bhinocricus, on the other hand, is abundant in South America and the West Indies, 
but not in North America. In Central America it extends from Panama to Mexico. 
It is also the dominant genus of the family Spirobolidae in the Oriental Region, but 
does not appear to extend in Eastern Asia so far north as Spirobolus, Clearly, there- 
fore, it is a more southern type than the latter. 

There is at present no agreement amongst systematists as to the number of families 
into which the Polydesmoidea should be divided, but, setting aside some of the peculiar 
forms referred to below, sufficient is known of the inter-relationship of some of the 
other genera to make instructive comparison between the Polydesmoid fauna of 
Central America and that of other countries. 

The most important of these belong to the families Platyrhachidae and Chelo- 
desmidae. 

The two subfamilies of Platyrhachidae are represented in the tropical south-eastern 
countries of Asia, the Platyrhachinee being especially abundant in Malaysia. They are 
not represented in tropical Africa. In America they inhabit the northern countries of 
South America, particularly Colombia and Ecuador, whence the Platyrhachina? extend 
northwards, at all events, to Costa Rica, and occur in the Antilles ; whereas the 
Euryurinse have a much more northern extension, passing from Costa Rica, through 
Guatemala and Mexico, into the Southern States of North America. 

The Chelodesmidse are very wide-ranging, but from our present point of view the 
most interesting forms are the species of the dominant Central American genus 
Bhysodesmus of the subfamily Xysiodesminae. This genus is represented by a vast 
number of species in the northern districts of Central America, but is apparently 
absent in South America and in the Antilles. It extends into North America, and 
closely allied, if not generically identical, forms occur also in China. 



DIPLOPODA AND PEOTOTfiACHEATA . 137 

The Central-American Diplopods, from the faunistic standpoint, belong to tliree 
categories: (1) genera with northern affinities and probably derived from the north; 
(2) genera with southern affinities and probably derived from the south ; (3) genera 
restricted or almost restricted to the country and probably autochthonous. 

The principal groups of northern origin are the Platydesmidae {Platydesmus), the 
Chordeumidae [Cleidogona), the luloidea (Paraiulus) ; Sjnrobolus of the group Spiro- 
boloidea and Rhysodesmus of the Xystodesmine Polydesmoidea. 

To the group with southern affinities and probably of southern origin must be 
referred the Spirostreptoidea {Orthoporus and Epinannolene), the Stemmiuloidea 
[Sfemmiidus) ; Bhinocricus of the group Spiroboloidea and several of the Poly- 
desmoidea, like Ligiodesmus, Oncodesmus (Oniscodesmidae), Lophodesmus (Pyrgo- 
desmidse), Platyrhachus, AmpUnus, and others (Platyrhachidae). Of these, Orthoporus 
and probably Epinannolene have African affinities; the Stemmiuloidea and Pyrgo- 
desmidse also have related genera in tropical Africa and Asia. On the other hand, the 
Oniscodesmidte and Platyrhachidae are at present known only from South-Eastern Asia 
and tropical America. The Oniscodesmidae may await discovery in Africa, but the 
Platyrhachidae appear to be certainly absent from that country. Their restriction to 
Indo- and Austro-Malaysia and tropical America is singular. Nevertheless, since their 
distribution agrees in a measure, though not in exact detail, with that of the existing 
Tapiroid mammals, it may be that they are tropical survivors of families once extending 
throughout the countries now bordering the North Pacific Ocean, 

Of the peculiar, probably autochthonous, groups, the most interesting are the 
Polydesmoidea of the family Sphaeriodesmidae, which appear to be a specialised offshoot 
of the Chelodesmidae. The family ranges in Central America from Mexico to Costa 
Eica, and one of the genera, Cydodesmus, is also represented in Jamaica. Also 
apparently restricted to Central America, with the exception of one form recorded from 
California, are the genera of the Chelodesmidae belonging to the subfamily Rhacho- 
desminae. 

Class PROTOTKACHEATA. 

No fossil forms of this class are known. On morphological grounds it is believed 
to be of great antiquity, older even than the Trilobites, but, as Gadow has pertinently 
remarked : — " This genus [Peripatus] need not date further back than the Cretaceous 
to allow us to account for the scattered distribution of its species " *. 

Until 1894 all the described species of this Class from South America, South Africa, 
and Australia, were referred to the single genus Peripatus, despite the ascertained 
existence of important morphological diiferences between those of the Continents in 
question. But since that date, when it was first proposed to give new generic names 

* Gadow hero ignores the work of Bouvier and others, who refer the Prototracheata to several genera. 
BIOL. CENTR.-AAiER., Introd. Vol., June 1915. T 



138 OEIGIN, ETC., OP THE FAUNA. 

to the South African and Australasian groups, and to restrict the original title to the 
South American groups, great additions have been made to our knowledge of this 
Class, mainly by the discovery of new genera, but partly by a more detailed acquaintance 
with species that had been previously described. One practical outcome of these lines 
of research has been rapid development of the classification from the point of view of 
nomenclature, with the result that there appears to be at least eight well-marked 
genera distributed as follows : — 

I. South and Central America and West Indies. 
Per'ipatus. Tropical America. 
Opisthopatus. Chile. 

II. Tropical and South Africa. 
Feripalus. Congo. 
Opisthopatus, South Africa. 
Peripatopsis. South Africa. 

III. East Indies. 

Typhloperipatus. Abor country, N.E. India. 
Eoperipatus, Sumatra, Malay Peninsula. 

IV. Australasia (including Australia, New Zealand, New Guinea, etc.). 

Paraperipatus. New Guinea, Solomon Islands, 
Peripaloides. Australia and New Zealand. 
Ooperipatus. Australia and New Zealand. 

With exception of Typhloperipatus, described in 1913, these genera were admitted 
by Bouvier in his Monograph of 1907-8. For my present purpose it is immaterial 
that one of the species referred by Bouvier to Ooperipatus has been recently given 
generic rank, under the name Symperipatus by Cockerel), and that the species from 
the Congo and Chile, assigned by the French author respectively to Peripatus and 
Opisthopatus, have been separated from those genera as Mesoperipatus and Meta- 
peripatus by Clark *. The interesting point in this connection is that close 
relationship exists, on the one hand, between the tropical African and tropical South 
American species and, on the other hand, between the Chilian and one of the South 
African forms. Clark's further proposals regarding the generic and subgeneric 
divisions, to which he refers the tropical American species assigned by Bouvier to 
Peripatus will be referred to later on. 

* It ■will be a nice controversial question for systematists to settle in the future whether the names 
" Congo-Peripatus " and " Chilio- Peripatus," proposed by Sedgwick for these same species, shall be admitted 
as generic terms or not. Clark, perliaps not wisely, disregarded them, probably because Sedgwick, true to his 
colours, expressly disclaimed the intention of putting them forward as generic terms. But since they 
were available as properly published names, future disputation would have been avoided by accepting 
them. 



PEOTOTEACHEATA. 139 

Three classifications of the genera into families and subfamilies have been put 
forward by Evans, Bouvier, and Clark. That of Clark, which is a compromise, with 
amplifications, between the other two, will serve to show the mutual relationships 
of the genera above mentioned admitted at the present time : — 

Family Peripatid^. 

Subfamily Peripatinse. 

Genus Peripatus {Mesopenpatus). Tropical America and Africa. 

Subfamily Eoperipatinae. 

Genus Eoperipatus. Sumatra, etc. 
Genus Typhloperipatus. N.E. India. 

Family Peripatopsidj!. 

Subfamily Peripatopsinse. 

Geuus Peripatopsis. S. Africa. 

Genus Paraperipatus. New Guinea, Solomon Islands. 

Subfamily Perii)atoidiua;. 

Genus Peripatoides. Australia and New Zealand. 
Genus Oopej-ipatus. Australia and New Zealand. 
Genus Opisthopatus {Metaperipatus) . Chile, S. Africa. 

Attention may be drawn to one or two interesting faunistic points revealed by this 
arrangement : — 

1. Resemblance, in the case of the Peripatidae, between the Malaysian forms and 

those of tropical West Africa and of tropical South America. This case is 
closely paralleled by that of the dwarf Squirrels, NanosciuridEe, amongst the 
Mammalia. 

2. The restriction of the genera of Peripatoidinae to Australia, New Zealand, 

Chile, and South Africa. 

3. The ascription of the Papuasian genus and one of the South African genera to 

the same subfamily, Peripatopsinae. These genera, however, are not very 
closely allied. 

The Central American Species. 

Bouvier pointed out in his Monograph that the tropical American species of 
Peripatus fall into two groups, which he named Peripates andicoles and Peripates 
caraihes. To the former, restricted to the mountains of Central America and the 
northern Andes, Cockerell gave the name Oroperipatus, limiting the term Peripatus 
to those found in the West Indies and lower-lying countries of South and Central 
America. This genus was subsequently divided by Clark into the following subgenera: — 
Plicatoperjpatus, known from one species {jamaicensis) from Jamaica; Macroperipatus, 

t2 



140 OEIGIN, ETC., OF THE FAUiS'A. 

containing species ranging from Eio Janeiro to Central America ; Epiperipatus, also 
containing many species with approximately the same distribution as Macropeiipatus ; 
and Peripatus (s. s.), restricted to species extending from Jamaica through the West 
Indies to Venezuela. 

Oroperipatus, occurring elsewhere in the Andes of Bolivia, Ecuador, and Colombia, 
is represented by two species in Central America, namely 0. eiseni from Tepic in 
Mexico and 0. goudoti from Mexico, without further particulars as to locality, 
although, as Bouvier suggested, it probably came from the western mountains of that 
country. 

Peripatus has many Central American representatives, namely perrieri, from Vera 
Cruz in Mexico, geayi, ranging from Cayenne to Panama, both belonging to the 
subgenus Macropen'patus ; biolleyi and isthnicola from Costa llica; nicaraguensis 
from Nicaragua; edwardsii, extending from Cayenne and Venezuela to Darien and 
Panama, and hrasiliensis, extending from Santarem on the Amazons to Panama, these 
five being referred to the subgenus Epiperipatus. 

It may be noted that Oroperipatus and Macroperipatus are absent from the 
Antilles, and that only one representative of Epiperipatus has hitherto been recorded 
from those islands, namely from Grenada, the southernmost of the chain *. 
On the other hand, Plicatoperipatus and Peripatus (s. s.) are absent from Central 
America ; while the latter subgenus, exemplified by peculiar species in Jamaica 
and Porto Kico and in most of the islands of the Lesser Antilles, exists also in 
Venezuela. 

Conclusions. 

The Prototracheata are at the present time restricted to tropical and south 
temperate latitudes. They have become adapted to widely different conditions 
so far as temperature and climate are concerned. Their extension in the Southern 
Hemisphere about as far as the 45th parallel (Tasm-ania, New Zealand, Isle of 
Chiloe), and their existence at high altitudes within the tropics precludes the belief 
that their apparent absence from southern temperate areas in the Northern Hemisphere 
is due to inability to maintain themselves to the north of the Tropic of Cancer or 
thereabouts. The evidence therefore is in favour of the view that the Class was 
southern in origin. 

The present distribution of the genera Peripatus {sensu lat.) and of Opidhopatus 
{sensu lat.) attests a former union between Africa and South America. The 
last-mentioned genus also attests, though le&s strongly, a union between Australasia 

* This instance recalls the occurrence of the Armadillo {Dasyimi) in Grenada, alone of all the 
Antilles. 



PJiOTOTEACHEATA. 141 

and South America, since its affinities appear to lie with the Australasian 
Feripatoides. 

In the case of the tropical American forms, it seems probable that Central America 
and the Antilles independently received their prototracheate faunas from South 
America. 

Recent Literatuke. 
BouviER, E. L.— Auu. Sci. Nat. Zool. (9) ii. pp. 1-376 (1905) * and v. pp. 01-318 (1907). 
Sedgwick, A.— Quart. Journ. Micr. Sci. lii. pp. 379-408 (1908). 

Clark, A. H.— Smith. Misc. Coll. l.'c. no. 17, pp. 1-5 (1913) j id. Proc. Biol. See. Wash. xxvi. 
pp. 15-20 (1913). 



* III justice to Mr. F. 0. Pickard Cambridge, ifc should be explained that in Bouvier's Monograph, figg. 1 
and 2, pi. i., depicting examples oiPeripatus simoni from Breves on the Amazons, are reproductions of water- 
colour sketches taken from life by Mr. Cambridge, who collected the specimens at that locality. The 
Bketchos were sent, with the specimens, to Bouvier from the British Museum, but apparently without any 
indication that Mr. Cambridge was the artist and collector. 



142 OEIGIN, ETC , OF THE FAUNA. 



Appendix. 

The recently published paper by W. D. Matthew, entitled "Climate and Evolution" 
(Ann. New York Acad. Sci. xxiv. pp. 171-318, Feb. 1916), came to hand after the 
Mammalian section of this work was in type and after the sections dealing with 
the Arthropoda were written. Two of the main propositions of this thesis have an 
important bearing upon suggestions that have been put forward to explain some of the 
facts in the distribution both of the Mammals and of the Arthropods : — 

1. "The principal lines of migration in later geological epochs have been radial 

from Holarctic centres of dispersal." (P. 172.) 

2. " The numerous hypothetical land-bridges in temperate, tropical, and southern 

regions, connecting continents now separated by deep oceans, which have 
been advocated by various authors, are improbable and unnecessary to 
explain geographic distribution. On the contrary, the known facts point 
distinctly to a general permanency of continental outlines during the later 
epochs of geologic time, provided that the allowance be made for the known 
or probable gaps in our knowledge." (P. 173.) 

As regards the Mammalia, mentioned in this part of the 'Biologia,' whose distribution 
is tentatively assigned to vanished land-bridges, Mr. Matthew holds the following 
opinions. The resemblance between Thylacinus and the extinct South American 
Sparassodont Marsupials, believed by some American palaeontologists to indicate close 
affinity, is held by Mr. Matthew to be adaptively convergent (p. 265) ; and Mr. Tate 
liegan has quite independently come to this conclusion (British Antarctic 'Terra 
Nova' Exped. 1910, Zoology, i. no. 1, pp. 41-43, 1914). 

Mr. Matthew thinks that the ancestors of the Australian Marsupials entered 
Australia by way of Southern Asia, while those of the South American forms came 
down from North America. Mr. Matthew ascribes the absence of fossil Marsupials 
from Asia to the " imperfection of the geological record," and does not discuss either 
their comparative paucity in the Austro-Malayan Islands or their absence to the 
west of Wallace's Line — facts, which in my opinion suggest evolution in Australia, 
followed by northward migration into Austro-Malaya. Since the Didelphyidae have 
survived in South America since the invasion of higher placental types, and have even 
successfully penetrated into North America, it seems to me that the disappearance of 
the Marsupials from Holarctica and their absence at the present time from Indo- 
Malaya can only be provisionally assigned to the unsuccessful competition with more 



APPENDIX. 143 

highly organised groups. However, since the hypothesis of the South Pacific land- 
bridge, so far as mammals are concerned, rests largely upon the alleged closeness of 
relationship between Thylacinus and, e. g., Prothylacinus, the question need not be 
further discussed until the debated point is settled, if capable of settlement, by the 
osteological experts concerned. 

The difficulty of the South American Monkeys Mr. Matthew also surmounts by 
appealing to convergent evolution (p. 216). That is to say, the resemblances between 
the Monkeys of the Old and New Worlds have been independently acquired from 
Asiatic and South American Lemurs respectively. 

This possibility was long ago considered, and rejected, by Mr. Beddard, but was 
adopted by Dr. Scharff. The supposed South American Lemurs — Mr. Matthew admits 
that this group is "very doubtfully represented in the early Tertiary formations of the 
Argentine " — must have passed, he thinks, into South America from North America 
during the Eocene, if I understand him aright. However that may be, I agree with 
Mr. Beddard that it is difficult to believe that the relationship between the Cebidae 
and Simiidse is no closer than Mr. Matthew suggests. 

In connection with the Hystricomorpha, Mr. Matthew admits that " we find serious 
difficulties " (p. 229). After discussing the question, he concludes: "I have been 
unable to frame any hypothesis which will fit all the facts of the distribution of this 

group, except by assuming that the South American Hystricomorpha reached 

South America from Africa in the Oligocene by over-sea raft-transportation. This 
involves so long a voyage that I hesitate to accept it as a reasonable probability, even 
though the winds and currents obviously favor transportation in this direction" 
(p. 231). Since this suggestion does not seem to be seriously entertained by its 
propounder, it does not call for further comment. 

The presence of Manatees in the tropical rivers of West Africa and tropical East 
America is explained by the hypothesis of the former existence of the genus up the 
eastern and western shores of the Atlantic into the Arctic Ocean, to a point where his 
map (p. 174) shows the extreme north-eastern corner of Greenland may have been 
connected with the north-eastern corner of Europe — that is to say, almost to the pole. 
This theory will certainly account for the facts ; but, until the genus Trichechits 
turns up iu far northern Tertiary deposits, there is no direct evidence to support it. 

Although Mr. Matthew deals mainly with the Vertebrates, he remarks in connection 
with the supposed transatlantic bridge between Africa and South America : " The 
supposed evidence in its favour from lower vertebrates and invertebrates is due, so far 
as I have been able to examine it, to a lack of appreciation of the principles of 
dispersal of races and of parallelism and of the imperfection of the geological record " 
(p. 231). Presumably he holds the same opinion regarding the supposed bridge 
between South America and Australia. Nevertheless, I cannot bring myself to believe 



144 OEIGIN, ETC., OF THE FAUNA. 

that the close likeness between the Australian Scorpion CercojjJionius and several 
South American genera nor the resemblances between the genera of trap-door Spiders 
referred to Migidse (p. 131) are due to parallel evolution. Nor is it at all intelligible 
to rae why the Scorpions and Spiders in question have disappeared entirely from all 
parts of the world except the southern continents, if at one time they were Holarctic 
in distribution. Nor can I think that raft-transportation will account for their present 
discontinuous range. Whether the land-bridge theory is the true explanation is, 
of course, quite another matter ; but it appears to me to be the least improbable of 
the four *. 

• Elevations of ocean-floors to connect continents are usually regarded as vast changes. From tlio human 
standpoint the epithet is justifiable; but, if the diameter of the earth be taken as a standard, such changes 
may be described as infinitesimal. 



[ 145 ] 



FLORA. 

By W. BoTTiNG IIemslet, LL.D., F.R.S, 

Since the publication of the Appendix and Introduction to the "Botany" of the 
' Biok)gia Centrali-Ameiica,' in 1888, botanical explorers have been very active, not 
only within our limits and in the contiguous countries, but also more especially in 
Africa and Eastern Asia, from Burma and China, southward and eastward, through the 
Malayan Archipelago to the Philippines, New Guinea, and Fiji. 

The addition of new species to the Central American Flora is enormous, especially 
from Mexico and Guatemala *. Of new genera, established on newly discovered 
types, there are relatively few ; but very many new genera have been founded by 
the segregation of old and familiar genera. 

Notable among the discoveries in tropical and subtropical districts are additional 
genera and species of southern types or families, belonging to the Vochysiaceae, 
Trigoniacese, Lecytbidacese, Lauraceae, Euphorbiaceae, Artocarpaceae, etc., etc. 

The identification of Schlechtendal's Mexican genus Jidiania, the discovery of 
several new species of this genus, and the founding of the family Julianiaceae, including 
the exceedingly rare and imperfectly known Peruvian raonotypical Orthopterygium, 
constitute a most interesting botanical contribution. 

But, taken on the whole, the internal discoveries throw no further light on phyto- 
geography than we possessed in 1888. So little was known of the natural history of 
Lower California that it was not included in the ' Biologia,' though politically and 
geographically belonging to Mexico. It has since been sufficiently explored, as well 
as the outlying islands of Guadalupe, the Revillagigedo Group, and others, to reveal 
the characteristics of the vegetation and flora, which are essentially Sonoran, with an 
intermixture in the north of Upper Culifornian species. The endemic element, alike 
in the Peninsula and the Islands, is of no greater proportion than that of the adjacent 
mainland. American botanists have devoted much time to the investigation of the 
^egetation of North Mexico and of the States and Territories north-west of Mexico 
with extremely interesting results, emphasising the distinct origin of the Pacific and 
Atlantic floras. 

* We have no statistics, except those kindly furaished for Guatemala by Captain John Donnell Smith, 
who, in his own publications, has added eighty-four genera and \'22\ species not included in the ' Biologia.' 
Eleven of the genera and 488 of the species are new. It may, however, be safely assumed that about 
2000 genera of flowering plants are represented in Central America by at least 15,000 species. — \V. B. H. 

siOL. CEXTR.-AMEK., Introd. Vol., June 1915. U 



14G OEIGIN, ETC., OF THE FLOEA. 

Outside of America, the exploration of Central Africa has yielded the most startling 
results. Contrary to expectation, the relationships between the African and American 
floras are almost as strong as those between the African and Asiatic floras, including 
representatives of such otherwise exclusively, or almost exclusively, American families 
as the Canellacese, Caricaceae, Humiriacete, Ilydnoracese, Loasacese, Mayacaceae, 
Napoleonaceae, Rapateaceae, Strelitziaceae, Velloziacese, and Vochysiacese. There are 
equally striking generic and specific connections. Altogether there are records of the 
same, or specially representative, genera and species belonging to upwards of sixty 
families or distinct groups. These facts, coupled with the zoo-geographical data 
and the palaeontology of the region, have given rise to the theory of a land-connection 
between Africa and South America in Eocene times, a theory accepted by both 
botanists and zoologists of high repute. Von Ihering's projected map of the assumed 
" Verteilnng von Land und Meer zur Eocan-Zeit " offers a plausible solution of some 
of the problems of the present longitudinal distribution of organisms, alike in southern 
latitudes and in remote islands. As long ago as 1885 (Introduction to the Botany of 
the ' Challenger ' Expedition), I rejected the theory of the vegetation of remote volcanic 
islands and groups of islands, such as the Galapagos and Hawaii, being necessarily 
of derived origin rather than remnants of a former wider flora. This view does not 
exclude dispersal by various agencies in distant parts of the world and natural 
migrations across the widest continents, as exemplified by some certainly introduced 
plants. 

The explorations of the last twenty years in Western and Central China furnish 
further evidence of the existence of close relationships between the floras of eastern 
temperate Asia and eastern North America. Types illustrating this feature are 
usually representative species of near affinity, as, for example, of the genera 
Liriodendron, Liquidambar, and Sassafras. This American-Asiatic element extends 
southward to Mexico, and is more prominent in the mountains of eastern South 
Mexico than it is in the north. The following genera of forest-trees represented in 
the Atlantic States, absent from the Pacific States, reach Mexico, namely: — Magnolia, 
Asimina, Tilia, Bohinia, Liquidamhar, Ilex, Liospyros, Bumelia, Ulmus, Celtis, Moms, 
Ostrya, Corpinus, and Carya. In a general sense, the western coast-forests are 
coniferous and the eastern deciduous. Certain genera are apparently now confined, 
or nearly so, to America and the Mediterranean region (including the Azores and 
Canaries) — such are Ilelianthemum, Lupinus, Heberdenia, Platanus, and Corema. 

Apart from the subantarctic flora, there are genera that extend from Chili to Mexico 
and from Australasia to Borneo. The magnoliaceous genus Drimys belongs to this 
category. Judging from analogies, this is an example of a northern extension of 
a southern type. The Proteaccoe, now almost exclusively confined to the Southern 
Hemisphere, furnish a similar instance in the closely allied genera Helicia and Roupala. 
The former ranges from Australasia to China, Japan, and India, the latter from 
Brazil to Mexico, and neither has further extensions. Of course, the absolutists 



BOTANY. 147 

would regard these as tardy stragglers in a southern migration. The phanerogamic 
flora of Central America (as calculated in 1888) comprised an intermixture of northern 
and southern types of varying proportions in different areas and an autochthonous 
element. The last constituted only eleven per cent, of the genera against seventy 
per cent, of the species. The percentages of genera restricted to America and of 
wider ranges were 53-7 and 4Q-3 respectively. For species the figures stand at 
89-9 per cent, restricted to America and 101 per cent, extending beyond America. 

The statistics of a specimen of the upper mountain-flora comprise 260 genera, of 
which eighty-two (or 31-6 per .cent.) were restricted to America, thirteen (or 5 per 
cent.) endemic within our limits, and the rest, 165 (or 634 per cent.), had extra- 
American extensions. 

Nine of the exclusively American genera had northern extensions ; twenty-five had 
southern extensions; twenty-two were common to the Andes only; and twenty-six 
were also represented in both North and South America. 

The total number of species enumerated is 604, whereof 504 (or 83*4 per cent.) were 
reckoned as endemic; 83 others (or 13'8 per cent.) not extending beyond America, 
with a residue of only 2'8 per cent, of wider range. The extensions to other parts of 
America are given as: N. America, 17; S.America, 19; Andes only, 39; N. and 
S. America, 8. Of course, most of these figures are rough approximations, but they 
are sufficient to show that the representatives of northern and southern types are 
nearly balanced, and that the Andine element in the mountain-flora of Central America 
preponderates over the temperate northern element *. 

Taking the whole phanerogamic flora of Central America, as it was known in 1888, 
638 of the genera had north-western extensions, 562 north-eastern connections, whilst 
1285 had southern extensions. Species yielded similar proportions. But statistics 
alone explain very little. 

The present complexities of plant-distribution point to greater and oftener repeated 
alterations in the distribution of land and water than is generally admitted, and no 
one theory is sufficient, in my estimation, to account for the origin and progressive 
dispersal of organisms. Indeed, it is doubtful whether sufficient evidence still exists 
to carry us to a convincing conclusion. There is always the great question whether 
organic as well as inorganic matter has not developed on the same or similar lines in 
different regions or centres. Comparing the following particulars of the distribution 
of families, it is difficult to realize that one part of the world has produced a land- 
vegetation wholly different in composition from that of any other part. Very diverse 
interminglings exist, but they give little or no clue to the beginnings. 

Accepting, for purposes of comparison, the number of families of flowering plants 
at 290, as defined in the seventh edition of Engler's ' Syllabus,' very nearly three- 
fourths are represented in both the Eastern and Western Hemispheres. In the more 

• Cf. Biol. Ceutr.-Am., liotany, iv. p. 299 (1887). 

U2 



148 OEIGIN, ETC., OF THE FLOltA. 

restricted sense of Bentham and Hooker, four-fifths are common to both Hemispheres. 
The families confined to America are twenty-five in number, namely, Batidacea;, 
Bromeliacese, Calyceracea;, Cannaceae (Scitaminese *), Caryocaracese (TernstrcEmiaceae), 
Columelliacese, Cyclantbacese, Cyrillaceae, Fouquieraceae (Tamariscinete), Gomortegaceae, 
Lacistemaceae, Leitneriaceae, Lennoaceae, Limnanthaceae (Geraniaceae), Malesherbiaceae, 
Marcgraviaceae (Ternstrcemiaceae), Martyniaceae (Peiialineae), Myzodendraceae, Qui- 
inaceae, San-aceniaceae, Thurniaceae, Tovariaceae (Capparidacese), and Tropaeolaceae 
(Geraniaceae). 

The foregoing families are of various categories. Eleven of these are limited 
to a single genus each, and the Bromeliaceae is the only one exhibiting a considerable 
development of genera and species, now numbering forty-five and about a thousand 
respectively, generally dispersed in tropical and south temperate America. Twelve 
out of these twenty-five families are not known to be represented within the limits 
of the ' Biologia.' 

Fourteen families are peculiar to the African region (including Madagascar and the 
Mascarene Islands), so far as known, eight of which are monogeneric. The only one 
of considerable development is the herbaceous Selaginaceae, associated with the 
Scrophulariaceae by Engler. The Chlaenaceae, comprising six genera and about 
twenty-five species, are peculiar to Madagascar. They belong to the Malvales, and 
are shrubs and small trees with showy flowers. 

A group of small and interesting families inhabiting eastern Asia, consisting of the 
Cercidiphyllaceae, Eucommiaceae, Pentaphylacaceae, Stachyuraceae, and Trochodendraceae, 
of remarkable affinities, is not represented in North America, but Engler places 
Pentaphylax near the American Cyrillaceae. 

In spite of its highly difiierentiated vegetation, the Australian flora counts few 
peculiar families ; indeed, the only ones absolutely limited to Australia are the 
Cephalotaceae, restricted to the singular Cephalotus follicularis, the Eupomatiaceae 
and the Tremandraceae, allied to the Pittosporaceae, which, except the widely spread 
genus Pittosporum, are all Australian, But there are some characteristic Australasian 
(including New Zealand, New Caledonia, etc.) families with a few solitary outliers — 
such are the Centrolepidaceae, represented by one species in China and one or two 
species in the extreme south of America, and the Epacridaceae, with a few outliers in 
Malesia and Polynesia, and the mouotypic Lebetanthus in Fuegia. The EucryphiaceiB 
{Eucryphia), limited to about four species in Australasia and Chili, and the somewhat 
numerous and generally dispersed Australian Stylidiacese are represented outside of 
Australasia by solitary outliers only in India, Malaya, and Fuegia. 

In addition to the above-named families, there are some twenty-five others peculiar 
to the Old World (with some extensions in Polynesia), but inhabiting two or more of 

* The family-name under which the groups in question appear in the " Botany " of the ' Biologia.' — W.B.H. 



BOTANY. 149 

the great divisions of the Eastern Hemisphere. No fewer than fourteen of these are 
monogeneric and inconspicuous in the vegetation, except Casuarina and Nepenthes. 
Casuarina and PandanacecS are prominent and widely spread in maritime districts, yet 
they have not reached America independently of human agency. The Flagellariaccae 
and Philydracea! also largely affect coastal regions. The arboreous Dipterocarpacete, 
numbering some 300 species, is tlie only large family in this category, and it 
forms a conspicuous feature in the forests of India and Malaya — rare in Africa, and 
apparently absent from Australia. 

Including the families respectively nearly or quite peculiar to Africa, to Eastern 
Asia, and to Australasia, there are fifty-three fiimilies in the Old World which are not 
represented in America, as against twenty-five restricted to the New AVorld. 

From the foregoing particulars of distribution it is evident that harmonies do not 
exist on a large scale in the same way that obtains, for example, in the different 
islands of the Galapagos Archipelago. The peculiarities of plant-distribution are 
infinite and reducible to no laws : take, for illustration, the composition and present 
distribution of the families of root-parasites in relation to their affinities — the widely 
spread Balanophoracea; and OrobanchaceiE and the local Lennoacea; of North Mexico, 
or such essentially insectivorous families as the Nepenthaceae of the Old World 
and the Sarraceniacea? of America and the widely spread Lentibulariacese. One 
might go on multiplying the exposition of the anomalies and curiosities or phenomena 
of distribution. But just one more example: the Lardizabalaceae comprise about 
half-a-dozen small genera, several of them distinct monotypes ; two of the genera are 
endemic in Chili, and the rest of the order is restricted to North India, Ciiina, and 
Japan. 

Mr. E. I. Pocock, in his account of the Mammalia, seems to have arrived at the 
same inconclusive results as myself in respect of the plants ; but he evidently favours 
a former southern land-connection as a solution of the main problem, and, barring 
independent developments of life, it is the only theory that commends itself to my 
mind. No other explanation seems adequate *. 

• It was originally planned to discuss more fully in this place the composition and origin of tho Central 
American Flora, and at least six months were devoted to the collection of materials for this purpose ; but a 
combination of adverse circumstances has hitherto hindered the completion of the work and rendered it 
impossible within the immediate future. It is a great disappointment to me that I could not furnish 
Dr. Godman with something in ray lino more worthy of his monumental publication. 

I have great pleasure in recording the fact that I have received much assistance from American Botanists, 
■who have most liberally presented their published works. My thanks are more especially due to Prof. T. 8. 
Brandegee, Dr. N. L. Uritton, Mr. H. Pittier, Dr. B. L. Kobinson, Dr. J. N. Rose, Captain J. Donnell Smith, 
and Dr. W. Trekase.— W. B. H. 



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