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The reason of this agreement, which they zealously guarded, was this: at their very first establishment in their several lordships an oracle was given them that that one of them who poured a libation from a bronze vessel in the temple of Hephaestus where, as in all the temples, it was their wont to assemble should be king of all Egypt. Though the pyramids were greater than words can tell, and each one of them a match for many great monuments built by Greeks, this maze surpasses even the pyramids.

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It has twelve roofed courts, with doors over against each other: six face the north and six the south, in two continuous lines, all within one outer wall. There are also double sets of chambers, three thousand altogether, fifteen hundred above and the same number under ground. We ourselves viewed those that are above ground, and speak of what we have seen; of the underground chambers we were only told; the Egyptian wardens would by no means show them, these being, they said, the burial vaults of the kings who first built this labyrinth, and of the sacred crocodiles.

Thus we can only speak from hearsay of the lower chambers; the upper we saw for ourselves, and they are creations greater than human. The outlets of the chambers and the mazy passages hither thither through the courts were an unending marvel to us as we passed from court to apartment and from apartment to colonnade, from colonnades again to more chambers and then into yet more courts. Hard by the corner where the labyrinth ends there stands a pyramid forty fathoms high, whereon great figures are carved. This lake has a circuit of three thousand six hundred furlongs, or sixty schoeni, which is as much as the whole seaboard of Egypt.

Its length is from north to south; the deepest part has a depth of fifty fathoms. That it has been dug out and made by men's hands the lake shows for itself; for almost in the middle of it stand two pyramids, so built that fifty fathoms of each are below and fifty above the water; atop of each is a colossal stone figure seated on a throne. Thus these pyramids are a hundred fathoms high; and a hundred fathoms equal a furlong of six hundred feet, the fathom measuring six feet or four cubits, the foot four spans and the cubit six spans. The water of the lake is not natural for the country here is exceeding waterless but brought by a channel from the Nile; six months it flows into the lake, and six back into the river.

For the six months that it flows from the lake, the daily take of fish brings a silver talent into the royal treasury, and twenty minae for each day of the flow into the lake. Sardanapallus king of Ninus had great wealth, which he kept in an underground treasury. Certain thieves were minded to carry it off; they reckoned their course and dug an underground way from their own house to the palace, carrying the earth taken out the dug passage at night to the Tigris , which runs past Ninus, till at length they accomplished their desire.

The Egyptians bore the earth dug out by them to the Nile, to be caught and scattered as was to be thought by the river. Thus is this lake said to have been dug. On the last day of the feast, they being about to pour libations, the high priest brought out the golden vessels which they commonly used for this; but he counted wrongly and gave the twelve only eleven. All the other kings too were wont to wear helmets, and were then helmeted; it was not in guile, then, that Psammetichus held out his headgear; but the rest marked Psammetichus' deed, and remembered the oracle which promised the sovereignty of all Egypt to whosoever should pour libation from a vessel of bronze; wherefore, though they deemed Psammetichus not to deserve death for they proved him and found that he had acted without intent , they resolved to strip him of the most of his power and chase him away into the marshes, and that he was not to concern himself with the rest of Egypt.

Therefore he held himself to have been outrageously dealt with by them and had a mind to be avenged on those who had expelled him, and he sent to inquire of the oracle of Leto in the town of Buto, which is the most infallible in Egypt; the oracle answered that he should have vengeance when he saw men of bronze coming from the sea. Psammetichus secretly disbelieved that men of bronze should come to aid him.

Psammetichus saw in this the fulfilment of the oracle; he made friends with the Ionians and Carians, and promised them great rewards if they would join him, and having won them, with the aid of such Egyptians as consented and these allies he deposed the eleven kings. Apis is in the Greek language Epaphus. Moreover he put Egyptian boys in their hands to be taught the Greek tongue; these, learning Greek, were the ancestors of the Egyptian interpreters.

The Ionians and Carians dwelt a long time in these places, which are near the sea, on the arm of the Nile called the Pelusian, a little way below the town of Bubastis. It comes of our intercourse with these settlers in Egypt who were the first men of alien speech to settle in that country that we Greeks have exact knowledge of the history of Egypt from the reign of Psammetichus onwards. There still remained till my time, in the places whence the Ionians and Carians were removed, the landing engines 24 of their ships and the ruins of their houses.

This Egyptian oracle is in a temple sacred to Leto, and is situated in a great city by the Sebennytic arm of the Nile, on the way upon from the sea. In Buto there is a temple of Apollo and Artemis. The shrine of Leto in which is the oracle is itself very great, and its outer court is ten fathoms high.

Another slab makes the surface of the roof, the cornice of which is four cubits broad. This lies in a deep and wide lake near to the temple at Buto, and the Egyptians say that it floats. However that be, there is a great shrine of Apollo thereon, and three altars stand there; many palm trees grow in the island, and other trees too, some yielding fruit and some not. The story told by the Egyptians to show why the island moves is this: when Typhon came seeking through the world for the son of Osiris, Leto, being one of the eight earliest gods, and dwelling in Buto where this oracle of hers is, received Apollo in charge from Isis and hid him for safety in this island which was before immovable but is now said to float.

It was from this and no other legend that Aeschylus son of Euphorion stole an imagination, which is in no other poet, that Artemis was the daughter of Demeter. For the aforesaid reason say the Egyptians the island was made to float. Such is the tale. It was he who began the making of the canal into the Red Sea, 25 which was finished by Darius the Persian.

This is four days' voyage in length, and it was dug wide enough for two triremes to move in it rowed abreast. It is fed by the Nile, and is carried from a little above Bubastis by the Arabian town of Patumus; it issues into the Red Sea. The beginning of the digging was in the part of the Egyptian plain which is nearest to Arabia; the mountains that extend to Memphis in which mountains are the stone quarries come close to this plain; the canal is led along the lower slope of these mountains in a long reach from west to east; passing then into a ravine it bears southward out of the hill country towards the Arabian Gulf.

Now the shortest and most direct passage from the northern to the southern or Red Sea is from the Casian promontory, which is the boundary between Egypt and Syria, to the Arabian Gulf, and this is a distance of one thousand furlongs, neither more nor less; this is the most direct way, but the canal is by much longer, inasmuch as it is more crooked. In Necos' reign a hundred and twenty thousand Egyptians perished in the digging of it. The Egyptians call all men of other languages barbarian.

He used these ships at need, and with his land army met and defeated the Syrians at Magdolus, 26 taking the great Syrian city of Cadytis 27 after the battle. He sent to Branchidae of Miletus and dedicated there to Apollo the garments in which he won these victories. Presently he died after a reign of sixteen years, and his son Psammis reigned in his stead.

When the Eleans came to Egypt and told the purpose of their coming, Psammis summoned an assembly of those who were said to be the wisest men in Egypt. These assembled, and inquired of the Eleans, who told them of the rules of the games which they must obey, and, having declared these, said they had come that if the Egyptians could invent any juster way they might learn this too.

The Egyptians consulted together, and then asked the Eleans if their own townsmen took part in the contests. Then the Egyptians said that this rule was wholly wide of justice: "For," said they, "it cannot be but that you will favour your own townsmen in the contest and deal unfairly by a stranger. Nay, if you will indeed make just rules and have therefore come to Egypt, you should admit only strangers to the contest, and not Eleans.

He was more fortunate than any former king save only his great-grandfather Psammetichus during his rule of twenty-five years, in which he sent an army against Sidon and did battle by sea with the king of Tyre. Apries sent a great host against Cyrene and suffered a great defeat. The Egyptians blamed him for this and rebelled against him; for they thought that Apries had knowingly sent his men to their doom, that by their so perishing he might be the safer in his rule over the rest of the Egyptians. Bitterly angered by this, those who returned home and the friends of the slain openly revolted.

And Amasis showed that this was not displeasing to him, for being made king by the rebel Egyptians he prepared to march against Apries. When Apries heard of it, he sent against Amasis an esteemed Egyptian named Patarbemis, one of his own court, charging him to take the rebel alive and bring him into his presence. Patarbemis came, and summoned Amasis, who lifted his leg with an unseemly gesture b being then on horseback and bade the messenger take that token back to Apries. When Apries saw him return without Amasis he gave him no chance to speak, but in his rage and fury bade cut off Patarbemis' ears and nose.

The rest of the Egyptians, who still favoured his cause, seeing the foul despite thus done to the man who was most esteemed among them, changed sides without more ado and delivered themselves over to Amasis. Apries' men marched against the Egyptians, and so did Amasis' men against the strangers; so they came both to Momemphis, where it was their purpose to prove each other's quality. So many classes there are, each named after its vocation. The warriors are divided into Kalasiries and Hermotubies, and they belong to the following provinces for all divisions in Egypt are made according to provinces.

None of these has learnt any common trade; they are free to follow arms alone. Thus much is certain, that this opinion, which is held by all Greeks and chiefly by the Lacedaemonians, is of foreign origin. It is in Corinth that artisans are held in least contempt.

This acre is a square of a hundred Egyptian cubits each way, the Egyptian cubit being equal to the Samian. These lands were set apart for all; it was never the same men who cultivated them, but each in turn. These men, besides their lands, received each a daily provision of five minae's weight of roast grain, two minae of beef, and four cups of wine.

These were the gifts received by each bodyguard. There he was sustained for a while in the palace, and well treated by Amasis. But presently the Egyptians complained that there was no justice in allowing one who was their own and their king's bitterest enemy to live; whereupon Amasis gave Apries up to them, and they strangled him and then buried him in the burial-place of his fathers.

This is in the temple of Athene, very near to the sanctuary, on the left of the entrance. The people of Sais buried within the temple precinct all kings who were natives of their province. The tomb of Amasis is farther from the sanctuary than the tomb of Apries and his ancestors; yet it also is within the temple court; it is a great colonnade of stone, richly adorned, the pillars whereof are wrought in the form of palm trees.

In this colonnade are two portals, and the place where the coffin lies is within their doors. Moreover great stone obelisks stand in the precinct; and there is a lake hard by, adorned with a stone margin and wrought to a complete circle; it is, as it seemed to me, of the bigness of the lake at Delos which they call the Round Pond. It was the daughters of Danaus who brought this rite out of Egypt and taught it to the Pelasgian women; afterwards, when the people of Peloponnesus were driven out by the Dorians, it was lost, except in so far as it was preserved by the Arcadians alone, the Peloponnesian nation that was not driven out but left in its home.

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Now at first he was contemned and held in but little regard by the Egyptians, as having been but a common man and of no high family: but presently he won them to him by being cunning and not arrogant. He had among his countless treasures a golden foot-bath, in which he and all those who feasted with him were ever wont to wash their feet. This he broke in pieces and made thereof a god's image, which he set in the most fitting place in the city; and the Egyptians came ever and anon to this image and held it in great reverence.

We would have you sit aloft on a throne of pride all day doing your business; thus would the Egyptians know that they have a great man for their ruler, and you would have the better name among them; but now your behaviour is nowise royal. Such too is the nature of men. Then those others, when he denied the charge that he had taken their possessions, would bring him to whatever place of divination was nearest them; and the oracles often declared him guilty and often acquitted him.

When he became king, he took no care of the shrines of the gods who had acquitted him of theft, nor gave them aught for maintenance, nor made it his practice to sacrifice there, for he deemed them to be worthless and their oracles to be false; but he tended with all care the gods who had declared his guilt, holding them to be gods in very truth and their oracles infallible.

These are the outer measurements of the chamber which is made of one block; its inner length is of eighteen cubits and four-fifths of a cubit, and its height of five cubits. Some again say that a man, one of them that heaved up the shrine, was crushed by it, and therefore it was not dragged within. It was Amasis, too, who built the great and most marvellous temple of Isis at Memphis. It was Amasis also who made the law that every Egyptian should yearly declare his means of livelihood to the ruler of his province, and, failing so to do or to prove that he had a just way of life, be punished with death.

Solon the Athenian got this law from Egypt and established it among his people; may they ever keep it! Of these the greatest and most famous and most visited precinct is that which is called the Hellenion, founded jointly by the Ionian cities of Chios, Teos, Phocaea, and Clazomenae, the Dorian cities of Rhodes, Cnidus, Halicarnassus , and Phaselis, and one Aeolian city, Mytilene.

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It is to these that the precinct belongs, and these are they that appoint wardens of the port; if any others claim rights therein they lay claim to that wherein they have no part or lot. The bloodlines are those of the royals, the Abhorsen, the Clayr, and the Wallmakers. After this, the entire Wallmaker line physically became the Great Charter Stones and the Wall that separates the Old Kingdom from Ancelstierre, to prevent contamination of their descent.

Under the influence of the Wall, magic, both Free and Charter, exists only in the Old Kingdom; but can be practised in northern Ancelstierre, and further south if there is a strong wind from the Old Kingdom. The Dead : The Dead are ghosts with both the inclination and the ability to resist the river of Death, who re-enter the world of Life. Though a rare few emerge into Life on their own power, most must be summoned by a necromancer or emerge near a broken Charter Stone where the Charter's influence has been severely diminished, creating a "door into Death" , or where many deaths have recently occurred.

All Dead are averse to running water, and most are unable to withstand direct sunlight. There are two classes of Dead: Lesser and Greater. The Greater Dead are usually represented by Dead from beyond the Fifth Gate spirits from the deeper realms of Death and correspondingly more powerful. Greater Dead, such as Fifth-Gate Resters or Dead Adepts, may exist in Life without a physical body making them much more difficult to destroy. Lesser Dead may be incapacitated by immersing them in running water or by destroying their physical bodies with Charter Magic or explosives; Shadow Hands are impossible to harm by strictly physical means, but may be unraveled by specialized Charter Magic spells or returned to Death by the necromantic bells.

Most Dead prey on the living to remain in Life. Constructs : Free Magic constructs are forms assumed by Free Magic elementals or powerful Dead spirits such as Kerrigor. Though such constructs may be destroyed, destroying the Elemental itself is much more difficult and typically the province of Free Magic.

Mordicants : These are fiery constructs of clay and blood, animated by Free Magic and guided by a Greater Dead spirit able to move between Life and Death at will. They are fierce combatants, and seem to have enhanced senses, able to track specific targets over hundreds of miles, resisting direct sunlight to do so. The word mordicant comes from a Latin term meaning gnawing. The most common elementals belong to specific "breeds" such as Stilken, Magrue, Jerreq, or Hish , while the most powerful are unique, or "of a singular nature". Though "many thousands" of Free Magic Elementals escaped the creation of the Charter, most were later imprisoned or enslaved by it.

Of the remainder, "no truly dangerous creature of Free Magic has woken in a thousand years, save to the sound of Mosrael and Saraneth, or by a direct summons using their secret names". Some cannot be destroyed except by a Free Magic sorcerer more powerful than they, or by immersion in running water though Free Magic creatures of the Third Kindred, or those infused with the essence of the Nine, are exempt from this rule.

Charter Magic is typically ineffective. Many may only act within a given function, which can be of indefinite complexity. Certain fixtures excepted such as a surcoat or insignia , Charter Sendings do not possess a concrete physical shape, and derive identity primarily from their function. They are capable of emotive response, but show little desire except to fulfill their mandates.

The bells of necromancy are seven eponymous bells used by necromancers to control the Dead, named after the Seven Bright Shiners who invested themselves in the Charter. From smallest to largest they are:. Kibeth , the Walker, which can give the Dead freedom of movement or force them to walk according to the ringer's intention;. Astarael , the Weeper, also named Sorrowful, which sends both ringer and auditor far into Death. Each bell has a specific power over the Dead and Free Magic creatures, and if used by a skilled necromancer, also on living people. An errant or improper ring can affect the caster instead of the target, or cause other adverse effects.

The Abhorsens' bells are a "free-willed blend of Charter and Free Magic", though the spells they cast are "pure Free Magic". The most advanced and powerful Free Magic elementals, of which seven created the Charter and are represented by the necromancers' bells. Of the remaining two, Yrael later became Mogget, the Abhorsens' companion; and Orannis, "last and mightiest of the Nine", opposed the Charter and was imprisoned by the Seven.

According to "An Extract of the Journal of Idrach the Lesser Necromancer", a text posted on the series' website, the correlation of the Seven and the necromantic bells includes the Precincts of Death, with each bell equalling a specific Precinct. This would suggest that the additional precincts are related to the 8th and 9th "Bright Shiners", although it is not known which Precinct corresponds to which Bright Shiner.

These Great Charters invested themselves entirely within the bloodlines and artifacts of the Old Kingdom, as opposed to Astarael and Kibeth, who retained enough of themselves to remain separate entities the Disreputable Dog points out that she is only Kibeth in a "hand-me-down sort of way". It is implied that Saraneth and Mosrael wove themselves into the Abhorsen and Clayr bloodlines respectively.

Dyrim is considered the Great Charter of the royal bloodline. Yrael, also known as Mogget, initially refused to take a side for or against Orannis, and was therefore later enslaved to the Abhorsen by the other immortals. Whenever unbound, he tries to kill the current Abhorsen; but, during Orannis' second binding, he assists in the binding ritual. To Sabriel, Lirael, and Sameth, Mogget appears as a small white cat; to Terciel, Sabriel's father and predecessor as Abhorsen, Mogget adopted a different unknown name and appeared as an albino dwarf.

Mogget cannot use his dwarf-form without the permission of the current Abhorsen or Abhorsen-in-Waiting: Jerizael, the forty-eighth Abhorsen, forbade him from doing so for reasons unknown. At the end of the series, he re-appears as a cat. In Lirael , when Orannis possesses Nicholas Sayre and speaks to the necromancer Hedge, the story of the Binding is told in song:. I'll sing you a song of the long ago. Seven shine the Shiners, oh! What did the Seven do way back when? Why, they wove the Charter then!

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Five for the warp, from beginning to end. Two for the woof, to make and mend. That's the Seven, but what of the Nine— What of the two that chose not to shine? The Eighth did hide, hide all away, But the Seven caught him and made him pay. The Ninth was strong and fought with might, But lone Orannis was put out of the light, Broken in two and buried under hill, Forever to lie there wishing us ill. Death consists of Nine Precincts divided by Nine Gates, through which a grey river flows. Almost everything in Death is a bleak grey, and a subtle grey fogginess limits visibility.

The river may also contain and conceal hostile dead beings, who attack living travelers. Dead spirits can cross only when aided by a Necromancer, or when the border is weakened by a concentration of often violent deaths. Dexterity and great willpower are required to resist the current, which is psychological as well as physical. Each gate responds to a Free Magic spell wielded by Abhorsens and Necromancers; Dead cannot pass any Gate unless they are very powerful. Each Precinct contains a different peril. The First Precinct is mostly knee-deep water, but has eddies and pools dangerous to the interloper.

Its Gate is a huge waterfall. Multiple landscape features have recently been shown to be important for the magnitude of gene flow and thus for explaining population genetic structure in amphibians. Roads, mountain ridges and open or dry habitats apparently reduce gene flow between populations [ 7 , 8 , 17 , 18 ] while forests, wetlands and rivers are associated with increased gene flow [ 7 , 19 ].

For example, in two neotropical species of the genus Craugastor , mountains and dry forests present ecologically unsuitable habitat and act as barriers to gene flow resulting in diverged genetic lineages [ 18 ]. Both genetic and ecological divergence can lead to reproductive isolation and speciation. Ecological speciation is the evolution of reproductive isolation as a result of divergent natural selection in different environments and can proceed in the presence or absence of gene flow [ 20 ]. Several general scenarios for speciation have been identified including genetic differentiation, geographic distribution and shift in ecological niches [ 21 ].

These scenarios associate speciation processes occurring in sympatry, parapatry or allopatry with presence or absence of ecological divergence. A combination of environmental niche models with phylogeographic analyses offers the possibility to explore the role of geographic and ecological separation for speciation between diverging lineages.

Although phylogenetic and phylogeographic information has been increasing recently, few studies have been conducted simultaneously on both fine and large scale population genetics in tropical anurans [ 13 , 22 , 23 ]. In the Neotropics, phylo-geographic studies have been mainly limited to craugastorid [ 13 , 18 ], strabomantid [ 24 , 25 ] and dendrobatid frogs [ 26 - 30 ]. These studies often revealed genetic lineages within species [ 23 , 26 , 27 , 30 ], rectified the taxonomic relationship between species or species groups [ 28 ], or uncovered previously unknown species diversity [ 24 ].

Examining ecological divergence in a phylogenetic context has been applied to only a very limited number of Neotropical frogs. These studies demonstrated climatic specialization along temperature and seasonality axes and identified important speciation mechanism for some clades [ 21 , 31 ].

For most Neotropical frogs, however, there is no information about their genetic population structure, genetic diversity within the species, presence of genetic lineages, or even species status and variability in ecological requirements. Still less is known about the relationship between genetic divergence, reproductive isolation and ecological divergence, and the relative importance of all these aspects for speciation. Allozymes analysis by Ryan et al. This was confirmed by Weigt et al. These results were further confirmed by a fine-scaled molecular study in an area of Costa Rica and Panama that encompassed both genetic lineages [ 34 ] and a phylogenetic study of the P.

The Northern and Southern lineages are separated by a distribution gap of about km in Central Costa Rica [ 40 ]. Map of study area. Map with sampling localities with genetic clusters identified by S tructure Figure 2, this study. The figure was modified from Figure 1 published in Ref. We first conduct a Bayesian analysis to detect population subdivision and zones of admixture based on highly polymorphic nuclear markers.

We further calculate population differentiation F ST and R ST between inferred populations and assign individuals to their most probable population of origin for estimating migration between populations. Our analyses include calculation of genetic diversity across the investigated area. We also estimate divergence time between the Northern and Southern lineage and between population clusters based on sequence divergence in the mitochondrial Cyt B gene. Then we integrate our parameter estimates obtained from classical population differentiation analyses F-Statistics and recently developed Bayesian statistics.

Furthermore we conduct environmental niche modeling based on climatic and altitude parameters to estimate the potential for ecological niche divergence between genetic lineages, to measure ecological landscape connectivity across the distribution gap and to predict the distribution of past and current suitable habitat. The allopatric distribution of the Northern and the Southern lineage suggests that genetic divergence is a result of geographic isolation. This evidence, however, is insufficient to support an allopatric model of divergence because distribution ranges could have been in contact in the past.

To explore this possibility, we project niche models to the last glacial maximum. A result showing equal or higher habitat unsuitability across the distribution gap during the last glacial would provide further support for the scenario of allopatric divergence.

We discuss the possibility that speciation has occurred between the Northern and Southern lineage based on the combined evidence of genetic and ecological data together with previously collected behavioral data. Moreover, this thorough population genetic and ecological analysis on a species for which the communication system, mating system and mate choice strategies are well studied will offer the opportunity to detect important areas such as hybrid zones, areas of low or high genetic diversity and areas that differ ecologically for studying new aspects of behavior in relation to the ecology and genetic composition of the population.

Afterwards Pr X K increased only insignificantly. Conducting the analysis separately for the Northern and Southern lineage resulted in the same geographic-genetic pattern, i. However the Southern lineage can be further subdivided in several genetically distinct population clusters. Posterior probability for population cluster. For higher Ks the likelihood is not or only very slightly increasing.

However there is still a relevant change in the Pr X I K by increasing K to 3 and 4, which relates to a further subdivision of the Southern lineage in several population clusters. The Northern lineage or cluster was located in North Costa Rica, i. The cluster adjacent in the north-west of the admixture zone ranged from the southern edge of the gap Ojochal through the lowland areas along the coast and the lowland areas in the Valle de General Buenos Aires and Valle de Cotobrus Potrero Grande. The cluster adjacent towards the south of the admixture zone ranged until El Forastero in Western Panama.

The sample locality on the Peninsula Osa was included in this cluster. As expected, the Northern lineage was most diverged from all other clusters. R ST values between the admixture zone and both adjacent clusters were non-significant. For pairwise R ST values see [Additional file 1 ]. Overall, the results point to higher gene flow among populations in the cluster North than in the southern clusters.

Migration rates per generation N m [ 5 ] among population clusters previously identified by S tructure , based on F ST below and R ST above diagonal values. These individuals are assumed first generation migrants. No migrants were detected between the Northern and Southern lineages. In general, allelic richness per population or cluster increased from the northern most localities towards South America. Also the two populations flanking the gap [La Junta 8 and Ojochal 9 ] retained lower allelic richness. Isolation by distance. Grey squares represent distances between one population from the North and one population from the South; White triangles represent distances between two populations from the North, and black diamonds represent distances between two populations from the South.

Genetic distance and time of divergence: The mean p distance uncorrected nucleotide difference between the Northern and Southern lineage was 0. Whereas all distances between the Northern and the Southern lineages were between 0. Estimated divergence times between North and South were between 4. North haplotype 1 , 2. Haplotype network. One sequence of haplotype 4 orange and one sequence of haplotype 9 red are from Piedras Blancas, the admixture zone in South Costa Rica as identified by S tructure. In fact, only one haplotype was found in the North. Cyt B diversities over all populations, the Northern and Southern lineage and population clusters.

The models generated by Maxent logistic output consisted of maps with logistic values LV ranging from 0 unsuitable habitat to 1 maximum suitability. The model for the Southern lineage predicted regions with high suitability in the Pacific moist forests of Provincia Puntarenas Costa Rica and western Panama. Under a logistic value LV threshold of 0. For the Northern lineage, the suitable range in Middle America, west from the gap, decreased from 1,, km 2 last glacial maximum to , km 2 current.

Niche modeling. Higher values indicate higher habitat suitability. See text for details. Connectivity and suitability across the distribution gap in Costa Rica: The analyses of suitability and connectivity suggested that the distribution gap in Costa Rica persisted during the last glacial maximum. The analysis of habitat connectivity suggested similar trends. For the Northern lineage, habitat resistance to reach the southern range was almost equal for current conditions and the last glacial maximum least-cost distance path was 6.

For the Southern lineage, resistance was much higher during the last glacial Taken together, these results suggest that geographic separation between genetic lineages persisted during glacial events. In general, the genetic pattern revealed by microsatellites is confirmed by sequences of the mitochondrial gene Cyt B.

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In contrast, gene flow between sample localities is higher in northern Costa Rican than in the southern population clusters. Isolation by distance explains a large amount of the genetic variation in both lineages; however, time of divergence between population clusters also plays a significant role for genetic differentiation. Environmental niche modeling revealed that the Northern and the Southern lineage differ significantly in habitat type. Suitable habitat for the Northern lineage is drier and occurs in pine-oak forests, while the Southern lineage is associated with moist forests.

Annual precipitation is significantly higher for the habitat of the Southern lineage. Since the last glacial maximum suitable habitat decreased for the Northern lineage but increased for the Southern lineage. The habitat connectivity across the distribution gap in central Costa Rica was lower during the last glacial maximum, suggesting that ecological factors have prevented secondary contact from at least the Pleistocene to the present. This result is consistent with the large divergence time and the absence of migration between lineages recovered by two independent analyses see below.

Thayer's Notes:

This indicates that the revealed population structure is fairly static given that mtDNA indicates ancient, historical events while microsatellites indicate more recent processes. In general, amphibian populations display a high level of spatial genetic structure, mainly when interpopulation distances exceed several kilometers [ 12 ]. Lampert et al. All studies, including this one, found a significant effect of isolation by distance, i.

Our Bayesian Structure analysis did not further subdivide the Northern lineage into subpopulations and the assignment and migration analyses points to high levels of gene flow between sample localities. Pairwise R ST values ranged from 0. Our results suggest that genetic drift causes genetic differentiation while subpopulations are genetically connected by migrating animals.

Marsh et al. On the other hand, Marsh et al. Since the distance between our sample localities is on average 30 km, the apparent gene flow at such distances is probably established by substantial movements among breeding ponds that are much closer to each other. In the North, the assignment of individuals to their population was lower than it was in southern populations [Additional file 2 ], where more individuals were assigned to their true locality. This indicates either higher gene flow in the North than among southern populations, or a population bottleneck followed by population expansion.

In North Costa Rica no mountains separate different sample localities. Nevertheless, gene flow between Palmar Norte, Cortez and Potrero Grande seems to be high while Buenos Aires appears to be more isolated. It is possible that gene flow between these three sample localities is conveyed by rivers.

Also, Osa 15 and Golfito 16 seem to be genetically more isolated within the southern populations then are the rest of the southern populations. Osa is more remote from other sample localities on the Peninsula Osa, while Golfito is located close to the coast and is separated from the lowland areas with the other sample localities by smaller mountains. Genetic assignment to the other sample localities is also lower in populations close to the distribution gap between La Junta 8 and Ojochal 9 indicating that gene flow and genetic diversity are reduced in peripheral populations. The calculation of divergence times based on COI and a mitochondrial mutation rate of 0.

Our calculations resulted in a divergence time of approximately 5 Myr between the Northern and Southern lineage which falls into the lower end of the confidence intervals calculated by Weigt et al. The difference between both studies in average divergence time estimates very likely rests upon the different geographical dimensions, i. While the Northern lineage occupies dry forests, the Southern lineage occurs in moist forests with higher annual precipitation.

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The low predicted suitability for the Southern lineage across the distribution gap in Costa Rica is consistent with the known species distribution. Suitability across the gap for the Northern lineage, however, was intermediate and within the range of known localities. Thus, the model suggests that conditions in a portion of the gap might be suitable for the Northern lineage. This incorrect prediction of the model could be the result of 1 the exclusion of an environmental variable that limits the species distribution range, 2 ecological interactions with other species across the gap e.

Because there are not conspicuous dispersal barriers in the region, we suspect that explanations 1 or 2 are the most likely. Niche modeling suggests that the absence of each lineage in the distribution gap correlates with ecological variables and therefore suggests that not only do the two lineages inhabit different habitats but there might be ecological divergence between the lineages. Niche conservatism is the tendency of species to retain aspects of their fundamental niche over longer time periods and limits the adaptation to ecological conditions at geographic barriers [ 45 ].

Our results show that the Northern and Southern lineages occupy different environmental envelopes and this suggests that their niches have been evolutionarily labile. The geographic separation was then maintained by niche conservatism and followed by niche specialization. The absence of the Southern lineage in the gap suggests that niche conservatism can promote allopatric speciation as proposed by Wiens and Graham [ 45 ].

At the moment our data cannot discriminate between both possibilities for lineage separation. In comparison with the present, suitable habitat during the last glacial maximum covered larger areas for the Northern lineage while it was more restricted in the Southern lineage. We assume that climatic changes from colder and drier climate to warmer and moister climate are responsible for this development.

Drier climate during the last glacial maximum also seems to explain the higher habitat resistance of the distribution gap for the Southern lineage. Since habitat resistance persists today it seems improbable that both groups will come into contact in the near future unless the current climate change will result in more favorable conditions across the gap.


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In this study we found that the lineages inhabit different habitats and thus suggests the possibility that they might be ecologically divergent. For Ecuadorian frogs of the genus Epipedobates , sister species with allopatric or parapatric distribution replace each other altitudinally or latitudinally emphasizing the role of environmental niche shift in speciation in neotropical frogs [ 21 ].

Ecological prezygotic isolation could arise as a result of adaptation to contrasting habitats or temporal isolation [ 46 ]. Adaptation to different habitats could entail behavioral reproductive isolation because signals involved in mate choice could be under ecologically based divergent selection. Examples include sexual preferences for ecologically selected body size and color pattern in fish or beak size in birds [ 20 ].

This possibility needs further exploration in our study species. Ron et al. The observed habitat differences are consistent with the recognition of each lineage as a separate species. Species level divergence is also suggested by some evidence of prezygotic isolation. However this was not true for northern females which produced nests and tadpoles with males from both lineages. The pattern of call divergence in the gap region in Costa Rica shows that several call variables tend to be more divergent near the gap.

This preference for the local call was stronger than that found in females from Gamboa, which is well within the southern group [ 47 ]. The results of this study offer the opportunity for new insights on the complex relationship between genetic distance, ecological divergence, call divergence and mate preferences. The areas between two genetic population clusters and areas adjacent to admixture zones are especially interesting for further behavioral observations. Such studies might determine whether there are abrupt changes in certain call parameters across genetically different populations, or how the female preference pattern varies across admixture zones.

Also, in the context of reproductive character displacement and reinforcement [ 49 , 50 ] coupled with measurement of gene flow, such areas provide exciting possibilities for future research prospects. Moreover the detected differences in ecology between the two main lineages should influence behavior. The northern frogs inhabit drier forests than the southern frogs. We would assume that temperature and precipitation pattern influence the time of the breeding period and the reproductive behavior.


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